THE  ESTABLISHMENT  OF  VARIETIES  IN  COLEUS  BY 
THE  SELECTION  OF  SOMATIC  VARIATIONS. 


BY 


A.  B.  STOUT, 

Director  of  the  Laboratories,  New  York  Botanical  Garden. 


WASHINGTON,  D.  C. 
Published  by  the  Carnegie  Institution  of  Washington 

1915 


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THE  ESTABLISHMENT  OF  VARIETIES  IN  COLEUS  BY 
THE  SELECTION  OF  SOMATIC  VARIATIONS. 


BY 


A.  B.  STOUT, 

Director  of  the  Laboratories,  New  York  Botanical  Garden. 


WASHINGTON,  D. 
Published  by  the  Carnegie  Instituti 

1915 


THIS  BOOK  IS  DUE  ON  THE  DATE 
INDICATED  BELOW  AND  IS  SUB- 
JECT TO  AN  OVERDUE  FINE  AS 
POSTED  AT  THE  CIRCULATION 
DESK. 


)00M/5-79 


Carnegie  Institution  of  Washington 
Publication  No.  218 


Copies  of  this  Book 
were  first  issued 

OCT  7    1915 


PRESS  OF  GIBSON  BROTHERS,  INC. 
WASHINGTON,  D.  C. 


TABLE  OF  CONTENTS. 


Page. 

Introduction  and  Historical  Review  of  Literature 3 

The  Problem  in  Coleus 12 

Method  of  Recording  Results 13 

Method  of  Handling  Cultures 16 

General  Survey  of  the  Variations 16 

Constancy  of  the  Various  Patterns 20 

Plants  with  yellow-red  blotched  pattern 21 

Plants  with  green-yellow-red  blotched  pattern 21 

Plants  with  color  pattern  green-yellow  spotted-red  blotched  and  with  uniformly  entire 

leaves 24 

Plants  with  laciniate  leaves 26 

Plants  with  green-red  blotched  pattern 29 

Plants  with  pattern  yellow-green-red  blotched 31 

Plants  of  color  pattern  green-yelloiv-solid  red 32 

Plants  with  green-solid  red  pattern 32 

Plants  with  green-yelloiv-solid  red  upper  center 33 

Plants  with  pattern  yellow-green-solid  red 34 

Plants  with  pattern  green-yelloio 34 

Plants  with  pattern  green  and  pattern  green-yellow  spotted 34 

Plants  with  pattern  yellow-green 35 

Range  of  the  Variations 37 

Frequency  of  the  Bud  Variations 39 

A.  Changes  involving  yellow  and  green 39 

I.  Increase  of  yellow  and  decrease  of  green. 
II.  Decrease  of  yellow  and  increase  of  green. 
III.  Reversal  of  the  relative  positions  of  green  and  yellow. 

B.  Changes  involving  the  epidermal  red 40 

I.  Increase  of  epidermal  red. 
II.  Decrease  of  epidermal  red. 

C.  Changes  involving  leaf-shape 41 

Summary  and  comparisons 41 

Distribution  of  the  Bud  Variations  among  Different  Clones 43 

Environmental  Influence 47 

Seed  Progeny 49 

History  of  Coleus 51 

Discussion .  .  .  , 58 

Summary ' 74 

Bibliography 77 

Explanation  of  Figures  in  Plates 79 

1 


THE  ESTABLISHMENT  OF  VARIETIES  IN  COLEUS  BY  THE 
SELECTION  OF  SOMATIC  VARIATIONS. 


By  a.  B.  Stout. 


INTRODUCTION  AND  HISTORICAL  REVIEW  OF  LITERATURE. 

Judgment  as  to  the  genetic  constitution  of  a  plant  is  based  on  the 
expression  of  characters  in  a  plant  itself,  in  members  of  a  selfed  progeny 
and  in  a  hybrid  progeny.  It  is  largely  through  the  study  of  the  last 
named  that  there  has  developed  the  conception  that  characters  are 
represented  in  cells  by  unit  factors.  In  considering  the  evidence  as  to 
whether  these  assumed  factors  are  in  any  sense  units,  it  is  highly 
essential  that  the  individual  be  studied  as  to  the  variability  of  char- 
acters and  the  range  of  expressions  exhibited  in  homologous  parts. 
For  this  purpose  so-called  bud  variations  are  especially  significant, 
since  they  represent,  perhaps,  the  extreme  of  spontaneous  somatic 
variability  and  suggest  that  quite  as  marked  variations  as  exist  in 
hybrid  progeny  may  develop  in  the  parts  of  a  single  individual. 

In  an  excellent  summary  of  all  cases  of  bud  variation  known  to  him, 
Darwin  (1868)  shows  that  the  phenomenon  is  widely  distributed  in 
the  plant  kingdom  and  that  it  may  affect  various  parts  of  a  plant.  He 
drew  the  highly  interesting  conclusions  that  they  include:  (1)  rever- 
sions to  characters  not  acquired  by  crossing,  (2)  reversions  in  hybrids 
to  parental  characters,  and  (3)  cases  of  spontaneous  variability. 
The  latter  he  considered  as  of  the  same  sort  as  appear  in  seed  progeny. 
According  to  Darwin's  views,  "long-continued  and  high  cultivation" 
are  conditions  that  induce  bud  variations,  but  he  recognized  that  cer- 
tain cases,  especially  those  when  only  single  buds  or  parts  of  buds  are 
changed,  do  not  seem  to  be  due  to  external  conditions.  Darwin  con- 
sidered in  general  that  bud  variations  are  evidences  of  the  almost 
unlimited  variability  that  plants  manifest,  due  to  the  nature  of  hving 
structures  and  the  exciting  causes  of  environment. 

Darwin  did  not  believe  in  fixed  hereditary  units.  He  held  that 
characters  may  respond  directly  to  the  effects  of  environment,  and  also 
exhibit  spontaneous  variability  both  in  seed  and  in  vegetative  propa- 
gation. Furthermore,  sexual  hybridization  was  considered  to  be  of 
influence  in  modifying  and  changing  inherited  characteristics. 

De  Vries  (1901,  vol.  i,  p.  39)  considers  that  bud  variations  are  spon- 
taneous changes  most  common  in  varieties  with  incompletely  fixed 
characters.  He  includes  (1889,  p.  13)  these  phenomena  under  the  head 
of  "dichogeny,"  a  general  conception  proposed  for  cases  in  which  the 
nature  of  the  organ  may  be  determined,  as  he  assumes,  by  external 

3 


ESTABLISHMENT    OF   VARIETIES   IN    COLEUS 


conditions,  but  he  also  assumes  that  the  internal  constitution  may 
admit  of  development  in  several  directions. 

De  Vries  discusses  bud  variation  with  special  reference  to  variega- 
tion. For  the  frequent  and  striking  cases  of  the  production  of  varie- 
gated branches  on  green  plants  and  the  development  of  green  branches 
on  variegated  plants,  he  offers  the  old  explanation  of  "latent  poten- 
tiality." They  belong  in  general  with  mutations  in  that  they  appear  as 
clear-cut  discontinuous  variations;  the  former  are  classed  as  progressive 
mutations  and  the  latter  retrogressive  (1901,  vol.  i,  p.  606). 

De  Vries  makes  a  most  comprehensive  analysis  of  the  nature  of 
variegation  and  concludes  (1901,  vol.  i,  p.  616)  that  the  capacity  for 
variegation  is  more  widely  distributed  in  the  plant  kingdom  as  a  latent 
or  semilatent  character  than  perhaps  any  other  character.  He  notes 
that  true  aurea  varieties  are  few  and  are  remarkably  constant.  Most 
variegated  races  show  rather  wide  fluctuations  and  constitute  what  he 
calls  intermediate  races.  His  scheme  (1901,  vol  i,  p.  424)  of  represent- 
ing the  relationship  is  as  follows : 


Normal 
character. 

Anomaly. 

I.  Original  species,  preen 

Active. 
Active. 

Semilatent. 
Latent. 

Latent. 
Semilatent. 

Active. 
Active. 

II.  Half  race,  rarelv  variegated 

III.  An  equilibrium  is  maintained. 

IV.  Eversporting  variety,  variegated  .... 
V.  Constant  varietv.  aurea 

In  a  later  pubhcation  (1913)  de  Vries  considers  that  the  pangens, 
which  he  assumes  are  directly  concerned  with  the  transmission  and 
expression  of  characters,  may  be  not  only  active,  semilatent,  and  latent, 
but  also  labile. 

The  transition  from  constant  green  varieties  through  variegated 
varieties  to  constant  aurea  varieties  is  conceived  to  be  dependent  on 
the  degree  of  activity  displayed  by  antagonistic  qualities  within  the 
cells.  The  essential  changes  are  conceived  to  be  intracellular  and  not 
dependent  on  qualitative  cell-divisions;  all  the  cells  are  potentially 
aUke,  but  different  processes  within  the  cells  give  differences  in  expres- 
sion. Thus  the  conception  of  de  Vries  does  not  consider  that  the  hered- 
itary physiological  units  (pangens)  are  fixed  and  uniform.  They  are 
subject  to  effects  of  environment  and  thej-  exhibit  spontaneous  change 
even  to  the  degree  of  sudden  appearance  bj'  progressive  mutation. 
All  these  phenomena  may  be  exhibited  either  in  vegetative  or  in  seed 
progeny.  Sexual  hybridization  may  entirely  modify  or  change  the 
effect  and  nature  of  the  basic  physiological  units  of  heredity.  De 
Vries's  general  attitude,  however,  places  the  emphasis  on  discontinuous 
or  mutational  changes  as  the  only  really  stable  variations. 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  5 

The  exhaustive  summary  of  facts  given  by  Cramer  (1907)  fully 
substantiates  the  views  of  Darwin  and  de  Vries  as  to  the  main  classes 
of  variegation  and  the  remarkable  gradation  in  degrees  of  constancy 
and  inconstancy  which  are  exhibited  by  the  various  types  and  by  their 
bud  variations.  Cramer  gives  the  most  complete  and  detailed  sum- 
mary of  the  known  cases  of  bud  variation  that  has  been  published. 
He  includes  (p.  18)  under  the  term  "  Knospenvariation"  all  cases  where 
a  character  suddenly  changes  in  a  plant  in  a  way  that  can  not  be  attrib- 
uted to  environmental  influence.  He  considers  that  there  are  three 
main  classes  of  bud  variation:  (1)  vegetative  segregations  in  hybrids; 
(2)  intermediate-race  bud  variations;  and  (3)  vegetative  mutations. 
Recessive  characters  in  hybrids  may,  he  assumes,  separate  out  by 
vegetative  cell-divisions,  and  dominant  characters  which  have  been 
latent  may  reappear.  His  conception  of  the  variability  of  characters 
in  an  intermediate  race  is  the  same  as  that  of  de  Vries.  Characters 
concerned  in  bud  variations  in  intermediate  races  show,  it  is  considered, 
great  irregularity  in  expression.  The  bud  variations  which  give  a  seed 
progeny  quite  constant  for  the  character  involved  are  classed  as  vege- 
tative mutations.  This  classification  emphasizes  the  fact  that  charac- 
ters involved  in  hybridization  may  come  into  expression  quite  irregu- 
larly, and  that  spontaneous  and  fluctuating  variations  are  common  in 
vegetative  development. 

Cramer  devotes  a  most  interesting  and  instructive  chapter  to  varie- 
gated plants,  showing,  especially,  their  wide  distribution  in  the  plant 
kingdom,  the  wide  fluctuation  in  the  degree  of  constancy  of  their  seed 
progeny,  the  range  from  vegetatively  constant  to  inconstant  types,  and 
the  frequency  of  bud  variations  in  variegated  plants.  He  considers  (pp. 
126, 127)  that  loss  of  variegation  can  occur  in  two  ways:  (1)  by  atavistic 
bud  variations  and  (2)  through  influence  of  external  conditions.  He 
notes  that  it  is  very  difficult  to  distinguish  fluctuating  variability  from 
mutation. 

The  general  variability  of  the  characters  concerned  with  variegation 
is  well  shown  in  his  discussion  of  changes  that  may  occur  in  a  single 
plant,  such  as  the  following:  In  apparently  inherited  types  of  variega- 
tion, the  seedlings  are  often  green  at  first.  Seedlings  may  have  at 
first  variegated  cotyledons,  then  a  few  green  leaves,  and  then  variegated 
leaves.  Some  biennial  plants  are  pure  green  during  the  first  year  of 
growth,  but  variegated  in  the  second  year.  Various  plants  exhibit 
a  periodicity  in  their  variegation,  being  green  in  spring  and  variegated 
later  in  the  summer,  or  vice  versa.  He  gives  numerous  cases  of  the 
appearance  of  variegation  by  bud  variation  on  green  plants  resulting 
in  new  varieties  or  in  the  duplication  of  types  already  known.  These 
exhibit  various  degrees  of  constancy. 

Various  adherents  of  Mendelian  doctrines  have  more  recently  dis- 
cussed the  transmission  of  characters  appearing  as  bud  variations, 


b  ESTABLISHMENT   OF   VARIETIES   IN   COLEUS 

attempting  to  explain  them,  as  they  do  all  heritable  variation,  by  the 
presence  or  absence  of  a  unit  factor.  While  this  is  the  general  attitude 
of  those  who  have  investigated  the  inheritance  of  variegation  and  the 
nature  of  bud  variations  involving  variegation,  the  results  of  their  studies, 
as  a  whole,  show  great  diversity  and  many  necessary  modifications  of 
the  general  Mendelian  doctrine  of  the  integrity  of  unit  characters  or 
unit  factors  and  the  purity  of  the  segregations  of  such  assumed  factors. 

East  (1908,  1910a)  considers  that  a  large  majority  of  the  known 
cases  of  bud  variation  are  due  to  the  loss  of  a  dominant  character  and 
that  70  per  cent  of  all  known  cases  are  color  variations.  His  discussion 
and  suggestions  do  not  claim  to  be  comprehensive  or  critical,  and  he 
excludes  from  his  treatment  all  cases  of  bud  variations  in  variegated 
plants,  because  some  types  of  these  are  known  to  be  pathological.  It 
is  interesting  to  note  that  he  states  that  no  important  potato  has  arisen 
as  a  bud  sport.  He  reports  four  cases  of  bud  variation  in  potatoes 
giving  white  from  red  or  pink  which  appear  to  be  constant;  also  several 
cases  giving  colored  or  purple  blotched  from  pure  white,  concerning 
which  no  data  are  given  except  the  statement  that  they  are  not  con- 
stant. East's  view  is  that  bud  variations  are  due  to  loss  or  latency  of 
hereditary  units  that  stand  for  characters. 

Bateson  (1909,  p.  273),  especially,  has  advocated,  on  theoretical 
grounds  largely,  that  bud  variations  are  due  to  qualitative  cell-divisions 
in  somatic  tissues,  giving  somatic  segregation  of  unit  factors.  The 
idea  is  quite  identical  in  its  main  features  with  Weismann's  concep- 
tion of  qualitative  divisions,  giving  tissue  differentiation  in  ontogeny. 

In  considering  the  nature  of  the  albomarginate  types  of  variegation, 
with  special  reference  to  the  origin  and  the  development  of  the  green 
and  white  areas,  Baur  (1909)  has  made  criticial  anatomical  studies  of 
a  white-margined  Pelargonium  zonale  which  indicate  very  clearly  the 
relationship  of  white  and  green  tissues  as  periclinal  chimeras  and 
explain  the  appearance  on  them  of  branches  wholly  green  or  white. 
In  testing,  by  crossing  experiments,  his  assumption  that  the  green  and 
white  tissues  are  each  pure,  he  is  forced  to  the  further  assumptions  that 
the  loss  of  green  in  this  case  is  due  solely  to  the  condition  of  the  chro- 
matophores  and  that  male  sex-cells  carry  chromatophores.  He  does 
not  consider  that  white  cells  can  arise  spontaneously  from  green,  or 
vice  versa.  Yet  this  probably  did  occur  when  the  chimera  was  first 
produced.  In  fact,  the  numerous  varieties  that  are  peripheral  chimeras 
not  only  in  Pelargonium  but  also  in  other  genera  indicate  that  such 
spontaneous  loss  of  power  to  produce  green  is  frequent.  We  may  say 
that  Baur's  interesting  results,  however,  do  show  that  if  spontaneous 
loss  occurs  in  young  leaves  after  they  are  formed,  mottled  or  striped 
variegation  is  produced,  but  if  the  loss  occurs  in  the  growing-point  itself, 
then  chimeras  will  result,  their  constancy  depending  largely  on  the 
relative  permanency  of  the  change. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  7 

Later,  Baur  (1910),  in  studies  of  seed  progeny  of  variegated  types 
that  appeared  spontaneously,  assumes  that  green  is  the  combined 
result  of  three  factors.  If  a  certain  one  (Z)  is  absent  the  tissue  is  color- 
less, if  Z  and  Y  only  are  present  then  a  chlorina  type  is  produced,  and 
if  N  is  present  with  Z  and  heterozygous  Y,  the  aurea  coloration  results. 
In  Antirrhinum  ma  jus  alhomaculatum,  Baur  found  8  types  of  variega- 
tion arising  spontaneously  in  cultures  of  about  30,000  pure-green 
plants.  His  crosses  with  these  show  that  certain  cases  appear  to  be 
inherited  only  from  the  seed  parent.  To  explain  the  results,  Baur 
assumes  that  hereditary  qualities  are  localized  in  different  parts  of  the 
cell.  The  nucleus,  the  cytoplasm,  and  the  chromatophores  all  possess, 
he  considers,  different  but  definite  factors  concerned  with  variegation. 

Such  results  and  conclusions  illustrate  very  well  the  difficulties  and 
uncertainties  which  arise  from  attempts  to  analyze  variations  in  terms 
of  unit  factors  and  suggest  most  forcibly  the  need  of  a  more  thorough 
investigation  of  such  variations  in  a  progeny  derived  by  vegetative 
propagation. 

The  variations  among  the  branches  of  a  single  plant  which  Correns 
(1909,  a  and  h)  reports  in  connection  with  variegated  types  of  Mirabilis 
jalapa  are  apparently  quite  similar,  in  degree  at  least,  to  those  I  shall 
report  for  Coleus.  It  would  seem  to  be  highly  important  that  the 
inheritance  of  these  variations  be  studied  in  vegetative  propagation. 
Correns,  however,  made  a  study  of  seed  progenies  only.  In  the  case 
of  the  albomaculata  type  these  were  composed  of  green,  white,  and 
albomaculata  plants  in  quite  different  ratios  for  different  plants  tested. 
All  these  classes  appeared  when  pollen  from  a  pure-green  plant  was 
used,  but  when  pollen  from  the  albomaculata  was  used  on  pure  green 
there  was  no  transmission  of  the  quality  of  variegation.  This  case  of 
matrocliny  is  due,  he  assumes,  first,  to  the  localization  in  the  cytoplasm 
of  the  factor  for  variegation,  and  second,  to  the  condition  that  male 
sex-cells  in  this  case  do  not  carry  cytoplasm. 

ShuU's  (1914)  studies  with  variegated  types  of  Melandrium  show 
much  the  same  results  as  those  of  Baur  and  Correns.  He  distinguishes 
between  chlorina,  pallida,  and  pure-green  types  of  Melandrium  on  the 
basis  of  presence  or  absence  of  three  factors  which  in  crosses  behaved 
quite  like  units.  In  types  with  green-white  blotched  and  with  chlori- 
nomaculata  variegation,  however,  the  variegation  seemed  to  be  trans- 
mitted only  through  the  seed  parent,  but  not  uniformly,  for  crosses  of 
variegated  branches  with  pure  green  gave  in  the  Fi  generation  plants 
ranging  from  pure  green  through  types  of  chlorinomaculata  to  yellowish- 
green  plants.  It  is  highly  interesting  that  Shull  found  that  the  greater 
the  amount  of  chlorina  coloration  in  the  calyx  the  greater  was  the  num- 
ber of  variegated  seed  progeny.  The  aurea  types,  which  possessed  as 
a  rule  small  round  flecks,  gave  such  varied  results  with  appearance  of 
different  types  in  Fi  progeny  that  Shull  concludes  it  must  be  an  infec- 


8  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

tious  variegation  which  is  transinitted  through  both  germ-cells  to  a 
part  of  the  progeny. 

Among  the  most  interesting  series  of  observations  especially  bearing 
on  the  behavior  of  red  coloration  in  Coleus  and  most  important  in  the 
consideration  of  the  nature  of  variegation  and  the  character  of  somatic 
variation  and  the  relations  of  these  to  seed  progeny,  is  that  of  Emerson 
(1914).  The  variegation  in  question  is  that  of  pericarp  color  in  certain 
"calico"  races  of  corn.  The  extent  of  coloration  varies  widely,  ranging 
from  solid  red  through  every  degree  of  striping  and  blotching  to  entirely 
non-red,  both  for  ears  as  a  whole  and  for  single  kernels  on  the  same  ear. 

Emerson  made  careful  studies  of  the  progeny  of  seeds  having  different 
degrees  of  coloration.  His  results  show  a  wide  range  of  variation  in 
the  progeny  of  kernels  that  appear  to  be  identical.  Selected  solid-red 
kernels  from  "freak  ears"  of  unknown  parentage  gave,  in  some  cases, 
progeny  with  only  solid-red  or  non-red  ears,  and  in  other  cases  the 
plants  produced  solid-red,  variegated,  or  non-red  ears.  Variegated  and 
white  kernels  (data  not  given  separately)  gave  either  variegated  and 
white,  or  red,  variegated,  and  white.  Again,  from  two  ears,  kernels  of 
white  gave  progeny  pure  white,  and  red  kernels  gave  red  and  white 
only,  each  of  which  gave  afterward  a  constant  progeny. 

In  selfed  variegated  strains,  kernels  of  all  classes  from  solid-red  to 
non-red  gave  progeny  with  ears  solid-red  or  variegated,  but  none 
with  no-red  ears. 

Data  are  given  collectively  for  progeny  of  5  solid-red  ears  (selfed). 
These  gave  solid-red  and  variegated  (p.  18).  Progenies  of  only  two 
plants  of  these  solid-reds  are  reported.  One  gave  again  plants  with 
solid-red  or  variegated  ears,  the  other  gave  only  solid  red.  The  num- 
bers grown  in  this  generation  were  respectively  9  and  16.  Emerson 
considers  from  these  data  that,  in  general,  red-eared  plants  behave, 
judged  by  progeny,  as  if  they  were  hj'brids  either  between  solid-red  and 
variegated  or  sohd-red  and  white  races. 

The  data  show  quite  clearly,  as  Emerson  points  out,  that  the  more 
red  there  is  in  the  seed  planted  the  larger  the  percentage  of  red  ears 
in  the  progeny.  The  variegated  race  is  therefore  far  from  constant. 
Selection  from  red  kernels  and  from  red  ears  give  a  strain  quite  constant 
for  solid-red,  but  Emerson's  data  are  far  from  conclusive  that  a  pure 
solid-red  strain  was  obtained  in  this  way. 

The  results  of  crossing  the  variegated  race  with  non-red  races  are 
interesting.  WTien  the  former  was  the  pollen  parent,  the  Fi  progeny 
gave  red,  variegated,  and  non-red  ears,  but  Emerson  states  that  some 
of  the  latter  may  have  been  extremely  light  types  of  variegation. 
From  the  reciprocal  cross  selected  kernels  for  the  Fi  gave  red  and 
variegated  or  only  variegated. 

In  the  selections  from  hybrid  stock,  as  in  that  in  selfed  stock,  the 
seed  that  had  more  red  gave  greater  numbers  of  pure-red  progeny. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  9 

Although  there  is  this  irregular  expression  of  solid-red,  variegated, 
and  non-red  kernels  on  the  same  plant  and  among  different  plants  of 
the  same  generation,  and  although  selection  from  solid-red  tends  to 
give  greater  numbers  of  solid-red  progeny,  Emerson  concludes  that  the 
factors  for  solid-red  and  variegation  are  "as  distinct  in  inheritance  as 
any  two  factors  could  well  be"  (p.  33).  He  points  out,  however,  that 
the  "factors"  concerned  are  pattern  factors,  one  determining  self- 
color  and  the  other  giving  variegation.  When  a  solid-red  kernel  occurs 
in  an  ear  of  the  variegated  race,  Emerson  assumes  that  a  V  factor 
changes  to  S;  but  when  these  red  kernels  do  not  give  solid-red  progeny, 
he  further  assumes  that  in  these  diploid  cells  one  of  the  pair  of  V  factors 
changes  to  S  and  that  such  somatic  changes  may  affect  an  area  of  cells 
including  macrospores. 

Taking  this  at  its  full  value,  we  note  that  as  the  occurrence  of  solid- 
red  kernels  is  frequent  the  hereditary  factor  V  is  fluctuating  and 
extremely  labile,  changing  to  S  readily.  For  V  and  S  to  be  distinct  in 
inheritance  under  such  conditions  is  hardly  conceivable,  for  they  are 
not  even  distinct  in  cell  lineage.  Emerson  thus  reflects  the  strong  ten- 
dency of  most  modern  Mendelian  investigators  to  regard  their  assumed 
factors  as  temporary  conditions  with  quite  fluctuating  activities. 

Such  conflicting  results  as  the  above,  obtained  even  by  careful 
pedigree  methods  of  study,  may  well  lead  one  to  question  whether  our 
kaiowledge  of  the  behavior  of  plant  characters  in  inheritance  and 
expression  has  advanced  much  beyond  the  views  of  Darwin. 

Such  studies  may  well  establish  the  general  breeding  values  of 
certain  characters  in  particular  cases  which  are  of  practical  significance. 
The  more  refined  methods  of  pedigree  have  shown  that  plants  with 
identical  appearances  may  give  quite  different  progeny,  and  that 
selection  for  relative  degrees  of  constancy  should  be  made  in  pedigreed 
lines  rather  than  in  mass  selection.  Mendelian  results  also  indicate 
the  possibilities  of  hybridization  followed  by  selection  in  pedigreed 
lines  of  hrybrid  progeny. 

In  its  theoretical  significance,  however,  Mendel's  original  work  has 
two  points  of  special  interest.  First,  it  embodied  the  conception  that 
all  structures  of  Hke  character  are  due  to  a  single  hereditary  unit;  for 
a  specific  example,  all  the  wrinkles  of  all  the  peas  on  a  plant  were  con- 
ceived as  represented  in  the  germ-cells  by  a  single  unit.  This  was  in 
decided  contrast  to  Weismann's  general  view  that  each  wrinkle  is 
represented  in  the  germ-cells.  The  second  point  of  special  interest 
pertains  to  the  behavior  of  these  assumed  units  in  the  formation  of 
reproductive  cells  and  their  behavior  in  fertilization.  Mendel  assumed 
that  they  segregated  as  units  independent  in  behavior  and  pure  in 
composition  from  the  unit  representing  the  contrasting  character.  By 
the  conception  of  the  purity  of  the  segregations  of  hereditary  units,  each 
representing  only  an  entire  and  complete  quality  or  character,  Mendel- 


10  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

ism  must  be  judged.  It  is  not  a  question  of  the  appearance  of  parental 
characters  in  hybrid  progeny,  but  of  the  purity  of  those  segregations. 

Mendel  himself  greatly  modified  the  conception,  gained  from  his 
studies  with  peas,  that  an  entire  quaUty  is  represented  by  a  single 
unit.  In  Phaseolus  he  crossed  P.  nanus  having  white  flowers  with 
P.  multifiorus  having  purple  flowers.  The  Fo  generation  of  31  plants 
had  flowers  ranging  from  white  to  pale  violet  and  purple  red  of  various 
grades.  Only  one  had  white  flowers  like  those  of  P.  nanus.  He  sug- 
gests that  the  color  of  flowers  (and  seeds  as  well)  in  P.  multifiorus 
*'is  a  combination  of  two  or  more  entirely  independent  colors"  (Mendel, 
p.  367).  To  Mendel,  therefore,  should  be  given  credit  for  the  con- 
ception of  multiple  factors,  later  developed  especially  by  Nilsson-Ehle 
(1909)  and  by  East  (19106). 

Very  soon  after  the  so-called  rediscovery  of  Mendel's  law  for  the 
behavior  of  certain  characters  of  Pisum  in  hybridization,  it  was  noted 
that  new  quahties  frequently  appear  in  the  F2  generation,  as  Mendel 
found  was  the  case  in  beans.  The  presence-and-absence  theory 
developed  by  Bateson  and  Punnett  (1905)  attempted  to  account  for 
this  increased  variabiUty  giving  the  appearance  of  new  types.  This 
gave  chance  for  dihybrid  ratios  from  what  was  apparently  a  simple 
pair  of  contrasting  characters,  the  "absences"  segregating  out  into 
expressed  characters  new  at  least  to  the  immediate  ancestry. 

The  next  important  modification  of  the  general  view  was  the  apph- 
cation  of  Mendel's  idea  of  what  we  now  call  multiple  factors.  Nilsson- 
Ehle  (1908  and  1909)  found  that  certain  crosses  in  cereals  gave  in  the 
F2  progeny  large  numbers  of  one  parental  type  and  relatively  few  of 
the  other  type.  For  example,  the  apparently  simple  cross  of  white- 
kerneled  wheat  (Predel)  with  a  red-kerneled  sort  (Swedish  Velvet 
Chaff)  gave  an  F2  progeny  of  about  63  red-kerneled  plants  to  1  white. 
Nilsson-Ehle's  explanation  assumes  that  the  red  character  in  the 
Swedish  Velvet  Chaff  is  due  to  three  independent  factors  which  are 
each  of  equal  value  and  that  any  one  can  produce  the  same  effect  as  all 
three.  We  note  that  the  variabiUty  of  the  F2  generation  was  not 
increased  over  that  of  parents,  but  that  the  ratio  showed  almost  com- 
plete appearance  of  one  character. 

East  (19105)  applied  the  term  to  the  same  sort  of  phenomena  as 
Mendel  did,  {.  e.,  to  increased  variabihty.  The  apparently  simple  cross 
of  white  with  j^ellow  corn  gave  in  the  F2  a  generation  with  few  white  but 
with  a  large  number  of  yellow  kernels.  The  latter,  which  were  of  all 
gradations  of  intensity,  were  grouped  into  two  classes  by  East. 

It  should  be  noted  that  such  a  group  of  ''factors"  having  among 
themselves  different  values  but  working  together  to  produce  a  single 
character  are  not  necessarily  independent  in  the  fertilizations  of  the 
variety  concerned.  Inside  the  variety  they  go  together.  When  certain 
crosses  are  made  they  separate.     This  is  but  another  waj'  of  saying 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  11 

that  the  hereditary  bearers  of  characters  appear  to  be  spht  up  and 
modified  by  crossing,  giving  in  some  cases  quite  new  characters. 

Thus  in  the  period  of  less  than  20  years  the  hterature  passes  from  the 
confident  treatment  of  characters  as  units  with  a  simple  shorthand 
representation  to  a  discussion  of  '^ factor s^^  that  under  some  conditions 
work  together  as  a  multiple  unit  and  under  other  conditions  separate, 
producing  equal  or  different  expressions.  The  assignment  of  different 
values  to  the  assumed  factors  as  diluters,  intensifiers,  inhibitors,  and 
the  conception  of  multiple  factors  that  can  separate  out,  giving  aber- 
rant ratios  or  new  expressions  and  even  almost  endless  intermediate 
gradations,  both  of  so-called  qualitative  and  quantitative  characters  all 
reduced  to  descriptive  terms,  add  nothing  to  the  fundamental  concep- 
tions of  Darwin  regarding  variability  in  hybridization.  The  extreme 
apphcation  of  the  multiple-factor  hypothesis  simply  means  that  small 
variations  are  inherited  equally  as  well  as  are  large  variations. 

The  greater  number  of  Mendehans,  mutationahsts,  and  adherents 
of  the  doctrines  of  pure  lines  seem  to  hold  that  the  unit  factors  are 
changeless.  Others  still  accept  Darwin's  general  views  of  the  modi- 
fiability  of  the  fundamental  units.  The  latter  view  is  especially  well 
developed  by  Castle  (1912),  who  states  (p.  356): 

"In  my  experience  every  unit-character  is  subject  to  quantitative  variation, 
that  is,  its  expression  in  the  body  varies,  and  it  is  clear  that  these  variations 
have  a  germinal  basis  because  they  are  inherited." 

Morgan  (1913)  considers  that  factors  are  labile  aggregates  subject 
to  rearrangement,  that  processes  of  mutation  and  reversion  are  reversi- 
ble, and  that  in  eversporting  varieties  mutation  and  reversion  are  regu- 
lar processes. 

Bateson  (1902,  p.  201),  in  a  defense  of  early  Mendelian  views,  makes 
the  following  admission: 

"We  have  to  consider  the  question  whether  the  purity  of  the  gametes  in 
respect  to  one  or  another  antagonistic  character  is  or  is  likely  to  be  in  the  case 
of  any  given  character  a  universal  truth.  The  answer  is  unquestionably  No, 
but  for  reasons  in  which  ancestry  plays  no  part." 

More  recently  (1914,  p.  322)  he  has  expressed  the  view  that  the 
conception  of  multiple  factors  is  in  his  mind  an  admission  that  there 
are  imperfect  segregations.     To  quote  further: 

"Segregation  is  somehow  effected  by  the  rhythms  of  cell-division,  if  such  an 
expression  be  permitted.  In  some  cases  the  whole  factor  is  so  easily  separated 
that  it  is  swept  out  at  once;  in  others  it  is  so  intermingled  that  gametes  of  all 
degrees  of  purity  may  result." 

Thus  it  seems  that  the  short-hand  system  of  representing  hypo- 
thetical germ-cell  units  is  often  not  only  cumbersome  but  inaccurate. 
Some  phases  of  it  may  be  useful  as  descriptive  terms,  but  the  method 


12  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

has  led  into  purely  speculative  fields  in  the  attempts  to  represent  and  ex- 
plain imperfect  segregations  and  variations  in  the  hereditary  qualities. 

Furthermore,  it  appears  in  final  analysis  that  the  extended  studies 
of  seed  progenies  have  not  contributed  anything  fundamentally  new  to 
the  knowledge  of  the  nature  of  plant  characters.  At  least,  the  analysis 
of  characters  in  terms  of  hereditary  units  has  failed. 

It  would  seem  that,  in  considering  these  problems,  the  studies  of 
variation  among  numbers  of  a  seed  progeny  is  no  more  important 
than  studies  of  variation  in  progenies  derived  by  vegetative  propaga- 
tion. The  latter  should  give  much  more  conclusive  data  regarding 
such  questions  as  the  constancy  of  characters  (or  of  assumed  factors), 
the  purity  of  apparent  segregations,  and  the  frequency,  constancy,  and 
nature  of  spontaneous  changes  in  the  expression  of  characters. 

It  is  evident  that  the  facts  regarding  bud  variation  involve  the  funda- 
mental questions  of  heredity.  When  such  variations  occur  in  a  plant 
that  can  be  propagated  vegetatively,  there  is  opportunity  to  apply  the 
pedigree  method  of  experimental  study  to  successive  generations  pro- 
duced by  vegetation  propagation.  The  nature,  frequency,  and  perma- 
nence of  such  changes  as  appear  can  be  studied  without  the  complica- 
tions that  are  associated  with  alternation  of  generations  and  fertilization 
as  they  normally  occur  even  in  selfed  seed  progeny.  Special  evidence 
regarding  the  "expression"  of  characters,  which  also  bears  on  the  ques- 
tion of  their  inheritance,  may  thus  be  obtained. 

THE  PROBLEM  IN  COLEUS. 

For  the  study  of  variation  along  the  lines  indicated  above,  I  have 
grown  a  series  of  833  plants,  all  descended  by  vegetative  propagation 
from  two  plants  of  a  variety  of  Coleus. 

Coleus  is  particularl)'  favorable  for  such  study  in  that  bud  variations 
are  frequent  and  the  plant  is  readily  propagated  vegetatively.  The 
leaves  are  in  pairs  which  alternate  on  a  square  stem,  making  but  four 
rows  of  leaves.  Bud  variations  that  appear  sectorially  can  thus  be 
traced  with  ease.  In  a  young  plant  lateral  branches  usually  start  to 
develop  from  the  axils  of  all  the  leaves  on  the  main  stem.  In  a  large, 
bushy  plant  many  lateral  buds  remain  dormant,  but  by  proper  pruning 
any  bud  can  be  forced  to  develop  or  it  can  be  propagated  as  a  cutting 
which  will  give  it  full  chance  for  development.  Large,  bushy  plants 
3  or  4  feet  tall  grown  out-of-doors  often  have  a  total  of  as  many  as 
300  branches.  In  the  greenhouse,  also,  plants  can  be  grown  from  cut- 
tings, with  the  production  of  many  leaves  and  branches.  The  ease  with 
which  the  plant  is  propagated  vegetatively  makes  it  possible  to  grow  a 
large  series  of  pedigreed  plants  from  any  cutting,  and  to  thus  test  the 
frequency  with  which  variations  appear,  the  constancy  of  the  different 
types,  and  the  purity  of  any  vegetative  segregations,  all  bearing  on  an 
analysis  of  the  nature  and  inheritance  of  the  characters  concerned. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  13 

METHOD  OF  RECORDING  RESULTS. 

To  record  fully  the  series  of  Coleus  plants  descended  through  vege- 
tative propagation  from  any  plant  or  any  particular  branch  used  as  a 
cutting,  the  pedigree  method  of  culture  has  been  used.  It  may  be  men- 
tioned that  the  application  of  the  pedigree-culture  method  to  plants 
propagated  vegetatively  is  much  simpler  than  its  use  in  seed  progenies, 
the  difficulties  of  which  have  been  ably  presented  by  Shull  (1908). 

Each  cutting  was  given  a  number  which  indicated  its  lineage.  The 
two  plants  serving  as  original  parents  were  numbered  1  and  3,  and  the 
first  digit  to  the  left  in  a  number  of  any  plant  indicates  the  original 
parent  plant.  For  the  progeny  of  plant  No.  1,  the  second  digit  indi- 
cates the  particular  main  lateral  branch  from  which  the  first  cuttings 
were  taken,  and  the  third  digit  is  the  number  of  the  particular  cutting. 
Additional  digits  indicate  successive  plants  grown  later  in  the  par- 
ticular line  of  descent.  Thus,  plant  121  (I  read  these  one-two-one, 
etc.),  122,  and  123  were  grown  from  three  cuttings  taken  from 
branch  No.  2  of  plant  No.  1.  Plants  1211,  1212,  and  1213  were  from 
cuttings  of  plant  121,  while  the  numbers  12111,  12121,  12131,  etc., 
were  given  to  plants  of  the  next  generation  of  cuttings.  Thus  the 
number  of  any  plant  gives  its  complete  pedigree  since  the  experiments 
were  begun.  Particular  data  regarding  the  cuttings  and  the  plants 
have  been  recorded  on  cards  and  filed  in  the  manner  of  a  card  cata- 
logue. This  enables  one  to  trace  readily  the  inheritance  of  any  varia- 
tion through  a  series  of  generations  and  to  compare  different  lines  of 
descent  from  any  point  in  the  culture. 

Following  the  suggestions  of  Webber  (1903)  and  Shull  (1912)  the  term 
''clone"  will  be  used  in  speaking  collectively  of  all  plants  descended  from 
any  one  plant  or  branch.  All  the  plants  derived  from  plant  No.  1 
constitute  a  main  clone,  itself  made  up  of  numerous  subclones.  The 
records  of  pedigree  enable  me  to  designate  these  as  clone  11,  clone  13, 
clone  117,  etc.  I  shall  use  the  term  "fine  of  descent"  to  include  the 
different  plants  that  were  the  parents  of  any  one  plant.  The  term 
"generation"  refers  to  the  plants  that  were  grown  during  the  same 
period. 

The  observations  here  reported  were  made  on  successive  generations 
of  pedigreed  plants  derived  by  vegetative  propagation  from  two  original 
plants.  These  parent  plants  were  alike  when  young  in  possessing  a 
color  pattern  that  can  be  characterized  as  a  mosaic  of  green,  yellow,  and 
red,  with  the  colors  distributed  as  shown  in  figure  2. 

In  referring  to  the  color  patterns,  it  seems  best  to  the  writer  to  use 
terms  that  are  sufficiently  descriptive  to  make  the  matter  concrete  and 
which  at  the  same  time  are  somewhat  compact.  The  color  of  the 
subepidermal  tissues  in  the  center  of  the  leaf  will  be  mentioned  first,  as 
green  or  yellow;  the  color  of  the  subepidermal  tissues  at  the  margin  will 
be  mentioned  next,  as  green  or  yellow,  and  last  the  character  of  the 


14  ESTABLISHMENT    OF   VARIETIES   IN    COLEUS 

epidermis  as  red  blotched  (both  surfaces),  solid  red  (both  surfaces),  or 
solid  red  upper  center.  In  these  compound  expressions,  hyphens  will 
be  used  to  separate  the  terms  descriptive  of  each  color  element.  In 
the  cases  with  colorless  epidermis  no  reference  is  made  to  this  condi- 
tion. The  colors  were  determined  by  Ridgway's  Color  Standards  and 
Color  Nomenclature.  The  color  pattern  of  the  parent  plants,  in  this 
paper  referred  to  as  a  green-yelloiv-red  blotched  pattern,  is  a  mosaic 
made  up  of  a  green  center  and  a  yellow  border  with  conspicuous 
epidermal  blotches  of  red.  The  yellow  is  amber  yellow  and  constitutes 
an  irregular  band  about  the  margin  of  the  leaf.  The  green  is  a  spinach 
green  and  is  chiefly  massed  in  the  center  of  the  leaf.  Over  the  green 
portions  the  red  appears  as  violet  carmine,  but  over  the  underlying  yellow 
areas  it  is  nopal  red.  The  three  color  elements  are  in  such  sharp  con- 
trast that  any  marked  variation  is  readily  noted.  Increase  or  loss  of 
either  the  yellow,  green,  or  red  is  conspicuous,  as  one  will  appreciate 
from  a  glance  at  the  plates  that  illustrate  this  article. 

Spontaneous  bud  variations  consisting  of  marked  alterations  in  color 
pattern  appear  either  in  single  leaves  or  groups  of  leaves,  or  in  single 
branches  or  groups  of  branches,  affecting  the  whole  or  a  part  of  the 
leaves  or  branches.  When  appearing  in  a  terminal  bud,  one  or  more 
leaves  have  a  pattern  differing  from  that  of  the  leaves  below.  When 
appearing  in  a  lateral  bud,  the  first  leaves  of  the  branch  possess  a 
pattern  different  from  that  of  the  subtending  leaf.  Those  appearing 
in  a  terminal  bud  have,  in  all  cases  observed  by  the  writer,  been  sec- 
torial in  the  main  branch  itself.  That  is,  the  change  has  appeared  first 
in  a  part  of  the  branch  only.  These  variations  carried  on  into  newly 
formed  branches  give  plants  bearing  two,  three,  or  even  four  distinct 
types  of  foliage,  with  differences  especially  marked  in  cases  of  single 
branches  with  sectorial  distribution  of  two  patterns.  The  rather 
simple  arrangement  of  the  leaves  and  branches  in  Coleus  enables  one  to 
trace  the  extent  of  a  variation. 

This  may  be  illustrated  by  a  variation  that  occurred  in  plant  No. 
1171.  In  this  variation  the  relative  positions  of  the  green  and  the 
yellow  became  reversed,  as  shown  in  the  two  leaves  reproduced  in 
figures  2  and  6.  When  the  cutting  was  made  in  April  1912,  all  the 
leaves  had  uniformly  green  centers.  On  one  of  the  first  pair  of  branches 
to  develop,  however,  all  of  the  leaves  had  the  yellow  in  the  center.  As 
further  branches  developed,  the  new  pattern  appeared  in  5  other 
branches.  The  plant  produced  13  pairs  of  branches  on  the  main  branch 
before  it  was  necessary  to  remove  the  terminal  bud  to  insure  proper 
development  of  the  lateral  branches.  All  of  these  branches  developed  at 
least  to  a  size  sufficient  to  show  the  color  pattern  of  the  leaves  borne. 
The  6  branches  with  the  new  pattern  were  contiguous  and  were  located 
as  indicated  in  diagram  1 .  The  plant  was  transplanted  to  a  large  pot 
and  kept  in  a  greenhouse  over  winter  and  then  grown  out  of  doors 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS. 


15 


during  the  summer  of  1913.  All  of  the  leaves  produced  by  the  6 
branches  in  question  in  the  17  months  of  growth  were  uniform  and 
constant  to  the  new  type  and  were  in  marked  contrast  to  the  fohage 
of  the  other  branches.  During  the  summer  of  1913,  two  bud  variations 
occurred  in  secondary  branches  in  the  upper  part  of  the  plant.  One 
was  a  sectorial  loss  of  yellow  giving  type  green-red  blotched  from  green- 
yellow-red  blotched  and  the  other  was  a  complete  loss  of  green  in  one 
branch  giving  type  yellow-red  blotched.  In  September  1913,  the  plant 
bore  four  distinct  kinds  of  color  pattern,  viz, 
yellow-red  blotched  (fig.  1),  green-yellow  red 
blotched  (fig.  2),  green-red  blotched  (fig.  5),  and 
yellow-green-red  blotched  (fig.  6).  The  bud 
variation  to  type  yellow-green-red  blotched  was 
sectorial  in  the  main  stem  for  a  vertical  dis- 
tance of  six  nodes,  but  was  not  complete  for 
the  entire  stem,  a  condition  shown  in  diagram  1 . 

The  greater  number  of  bud  variations  first 
appeared  in  single  lateral  branches  and  not  in  a 
series  of  branches  on  a  main  stem,  as  described 
above  for  1171.  Where  such  a  variation  was 
sectorial  in  a  branch  the  continued  growth  gave 
more  or  less  irregular  extension  of  the  new  type. 

The  parent  plant  here  designated  as  No.  1 
was  one  of  several  Coleus  plants  which  were 
grown  at  the  New  York  Botanical  Garden 
during  the  sununer  of  1911.  This  plant  pos- 
sessed in  September  1911,  when  first  observed 
by  the  writer,  two  branches  bearing  leaves  in 
which  the  yellow  was  apparently  almost 
entirely  absent.  These  two  branches  were  in  the  same  rank,  one 
directly  above  the  other.  x\bout  one-third  of  the  entire  foliage  of  the 
plant  was  borne  by  these  two  branches  and  the  marked  green  aspect 
of  this  part  of  the  plant  was  in  decided  contrast  to  the  conspicuous 
yellow  in  the  foliage  of  the  rest  of  the  plant.  Upon  careful  examina- 
tion, a  few  yellow  spots  could  be  seen  in  many  of  the  leaves  of  one  of 
the  green  branches  (branch  14)  quite  like  those  of  the  leaf  shown  in 
figure  4.  The  leaves  on  the  other  green  branch  (branch  13)  were 
apparently  free  from  all  yellow  areas  (fig.  5). 

The  decided  loss  of  yellow  in  these  two  branches  constituted  the 
only  variation  in  the  dozen  or  more  plants  in  this  particular  bed  of 
Coleus.  To  test  the  constancy  of  this  variation,  as  well  as  the  reap- 
pearance of  it  and  of  other  variations,  pedigreed  cuttings  were  made 
from  each  of  four  main  lateral  branches  of  the  plant. 

About  the  same  time  random  cuttings  were  made  from  the  bed  of 
plants  for  stock  for  general  planting.     One  of  these  cuttings  produced 


2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

2 

6 

2 

(j 

2 

6 

2 

2 

6 

6 

2 

2 

6 

Diagram  1. — Position  of  the 
six  branches  on  plant 
1171  having  color-pattern 
ycllow-green-red  blotched 
(6)  among  those  ha\ang 
green-yellow-red  blotched 
patterns  (2). 


16  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

during  the  winter  of  1911-12  a  single  branch  bearing  leaves  with  the 
red  completely  covering  both  surfaces.  This  color  pattern  designated 
as  green-yelloiv-solid  red  is  shown  in  figure  8.  All  the  leaves  on  the 
branch  were  uniform  for  this  pattern  and  were  in  most  conspicuous 
contrast  to  the  rest  of  the  plant.  From  this  plant  a  number  of  pedi- 
greed cuttings  were  also  made. 

METHOD  OF  HANDLING  CULTURES. 

The  first  generation  of  plants  (series  111,  121,  131,  141)  was  grown 
in  a  greenhouse  during  the  winter  of  1911-12.  In  April  1912,  cuttings 
(series  1111,  1211,  etc.)  were  made  from  these.  During  the  summer  all 
of  the  plants,  both  old  and  young,  were  grown  out  of  doors  in  beds. 
In  the  autumn  cuttings  were  again  made.  The  plants  developed 
from  cuttings  taken  in  the  autumn  were  under  observation  for  a  year, 
7  months  of  which  they  were  grown  under  greenhouse  conditions. 
Cuttings  taken  in  spring  were  grown  only  out  of  doors.  This  method 
of  handling  gave  opportunity  to  observe  development  and  behavior 
under  different  conditions  and  to  compare  old  plants  with  j^ounger 
ones.  Except  for  a  few  plants  that  were  subjected  to  special  condi- 
tions, all  the  plants  of  any  generation  were  treated  uniformly  with 
respect  to  kind  of  soil,  size  of  pots,  and  conditions  of  temperature 
and  illumination.  The  plants  were  cut  back  somewhat  to  prevent 
early  blossoming  and  to  maintain  a  vigorous  vegetative  condition. 

In  the  period  of  three  years  between  September  1911  and  September 
1914,  a  total  of  833  plants  were  grown  to  maturity  and  discarded.  All 
of  these  descended  through  vegetative  propagation  from  plants  Nos.  1 
and  3,  both  of  which  had  originally  the  green-yellow-red  blotched  pattern 
illustrated  by  figure  2. 

GENERAL  SURVEY  OF  THE  VARIATIONS. 

Variations  in  the  color  patterns  of  the  plants  both  of  the  original 
and  the  derived  types  can  be  classed  as  fluctuating  variations  and  as 
bud  variations.  In  the  former  the  changes  were  usually  quite  gradual 
and  affected  in  most  cases  an  entire  plant.  The  changes  which  are 
in  this  paper  included  in  the  term  ''bud  variations"  were  those  affecting 
only  a  part  of  a  plant  and  usually  appearing  as  a  sudden  and  conspicu- 
ous change.  In  addition  to  the  variations  in  color  patterns,  there 
appeared,  in  several  subclones,  plants  which  fluctuated  in  leaf-shape, 
giving  in  extreme  cases  leaves  deeply  cut  and  laciniate.  The  variations 
that  appeared  were  as  follows:  (A)  changes  involving  yellow  and 
green;  (B)  changes  involving  the  epidermal  red,  and  (C)  changes  involv- 
ing leaf-shape.  The  bud  variations  can  be  grouped  as  in  table  1  with 
data  regarding  the  number  of  plants  concerned  and  the  number  of 
times  the  different  changes  appeared  as  a  bud  variation. 


North  Carolina  Sti^Library 
.    i^teigh 


<:■ 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS. 


17 


Table  1. — Types  and  freqiiency  of  Ihe  bud  variations. 


Types  of  bud  variation. 


a 
a    . 

1^  o 


3 


c 


2. 


3. 

4. 


II. 


B. 


Changes  involving  yellow  and  green: 

I.  Increase  of  yellow  and  decrease  of  green: 

1.    Yellow-red  blotched  (fig.  1)  from  green-yellow-red  blotched  (fig.  2) 

Yellow-red  blotched  (fig.  1)  from  green-yellow  spotted-red  blotched  (fig.  4).. 

Yellow-red  blotched  (fig.  1)  from  yellow-green-red  blotched  (fig.  6) 

Yellow-solid  red  from  green-yellow-solid  red  (fig.  8) 

Green-yellow-red  blotched  (fig.  2)  horn  green-yellow  spotted-red  blotched  (fig.  4) 
Green-yellow-red  blotched  (fig.  2)   from  laciniate,  green-yellow  spotted-red 

blotched 

Green-yellow  spotted-red  blotched  (fig.  4)  from  green-red  blotched  (fig.  5) .  .  .  . 

Spontaneous  yellow  from  green-red  blotched  (fig.  5) 

Decrease  of  yellow  and  increase  of  green: 

1.  Green-yellow  spotted-red  blotched  (fig.  4)  from  green-yellow-red  blotched  (fig.  2) 
Green-yellow  spotted-red  blotched  (fig.  4)  from  yellow-green-red  blotched  (fig.  6). 

2.  Green-red  blotched  (fig.  5)  from  green-yellow-red  blotched  (fig.  2) 

Green-red  blotched  (fig.  5)  from  green-yellow  spotted-red  blotched  (fig.  4) .  .  . 
Green-red  blotched  (fig.  5)  from  laciniate,  green-yellow  spotted-red  blotched . . 

Green-red  blotched  (fig.  5)  from  yellow-green-red  blotched  (fig.  6) 

Green-solid  red  (fig.  9)  from  green-yellow-solid  red  (fig.  8) 

Reversal  of  position  of  green  and  yellow: 
1.   From  yellow  border  to  yellow  center 

a.  Yellow-green-red  blotched  (fig.  6)  from  green-yellow-red  blotched  (fig.  2) 

b.  Yellow-green-solid  red  (fig.  11)   from  green-yellow-solid  red  (fig.  8)  .  . 
Changes  invohnng  the  epidermal  red: 

I.  Increase  of  red: 

1.  Green-yellow-solid  red  (fig.  8)  from  green-yellow-red  blotched  (fig.  2) 

2.  Green-yellow  spotted-solid  red  from  green-yellow  spotted-red  blotched  (fig.  4) . 

3.  Green-solid  red  (fig.  9)  from  green-red  blotched  (fig.  5) 

Decrease  of  red : 


III. 


II. 


1. 
2. 
3. 


Green-yellow  spotted  from  Uiciniate,  green-yellow  spotted-red  blotched. .  . . 

Green-yellow  spotted  from  green-yellow  spotted-red  blotched  (fig.  4) 

Green-yellow  (fig.  12)  from  green-yellow-red  blotched  (fig.  2) 

Green-yellow  (fig.  12)  from  green-yellow-solid  red  (fig.  8) 

Green  (fig.  13)  from  green-red  blotched  (fig.  5) 

Green  (fig.  13)  from  green-solid  red  (fig.  9) 

Yellow-green  (fig.  14)  from  yellow-green-red  blotched  (fig.  6) 

Decrease  of  red  with  concentration  in  epidermis  of  upper  surface: 

1.  Green-yellow-solid  red  upper  center  (fig.  10)  from  green-yellow-solid  red. 
C.   Changes  involving  leaf-shape: 

1 .  From  entire  to  periodically  laciniate 

2.  From  periodically  laciniate  to  constantly  entire 


5. 


III. 


337 

198 

41 

54 

198 

68 
90 
90 

337 
41 
337 
198 
68 
41 
54 


337 
54 


337 

198 

90 

68 
198 
337 

54 

90 
8 

41 

54 

765 
68 


7 
1 
3 
1 
3 

1 
2 
9 

8 
1 

24 
2 
1 

10 
4 


6 
2 


4 
2 
2 

1 
2 
10 
1 
3 
1 
1 


13 
1 


The  names  given  to  the  different  patterns  embody  the  principal 
features  of  coloration  on  the  basis  explained  above  (see  pp.  13  and  14). 
The  patterns  selected  are  with  one  exception  those  that  appeared  as  con- 
spicuous bud  variations  and  which  are  sufficiently  distinct  for  ready 
identification.  Numerous  other  types  that  are  intermediate  between 
the  types  given  could  also  be  designated  by  still  more  exact  classifica- 
tion.    The  following  descriptions,  together  with  the  colored  plates 


18  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

illustrating  the  types  as  classified,  will  enable  the  reader  to  visualize 
the  patterns  referred  to  by  name. 

Color  pattern  yellow-red  blotched  (fig.  1):  Leaves  almost  entirely 
amber  yellow  with  only  very  limited  and  scattered  areas  of  greenish 
tissue.  The  island-like  areas  of  green  are  surrounded  by  yellow. 
Irregular-shaped  blotches  of  nopal  red  are  scattered  over  both  upper 
and  lower  surfaces.  This  decidedly  yellow  pattern  was  derived  from 
the  several  patterns,  as  shown  in  table  1,  by  a  sudden  and  a  conspicu- 
ous loss  of  green  tissue. 

Color  pattern  green-yelloiv-red  blotched  (fig.  2) :  This  is  the  pattern 
possessed  originally  by  the  two  parents  and  has  already  been  described. 

Color  pattern  green-yellow  spotted-red  blotched  (fig.  4) :  In  this  pattern 
there  is  no  definite  border  of  yellow.  The  yellow  appears  in  rather 
limited  and  somewhat  scattered  areas,  sometimes  nearly  limited  to  the 
border  zone,  but  often  quite  generally  distributed  throughout  the  leaf. 
The  pattern  is,  therefore,  decidedly  greener  in  appearance  than  that 
of  the  parental  type. 

Color  pattern  green-red  blotched  (fig.  5) :  This  is  a  bicolored  pattern 
of  green  and  red.  As  there  is  no  underlying  yellow  the  epidermal  red 
appears  uniformly  as  violet  carmine.  This  t^-^pe  arose  frequently  on 
plants  with  patterns  containing  yellow  by  what  was  apparently  a  com- 
plete loss  of  yellow. 

It  may  be  noted  that  in  the  four  patterns  as  arranged  above  there 
is  an  increase  of  green  and  a  corresponding  decrease  of  yellow,  with 
the  distribution  of  the  epidermal  red  quite  uniform.  The  yellow-red 
blotched  pattern  gives  the  extreme  development  of  j^llow  with  almost 
complete  absence  of  green.  The  green-red  blotched  pattern  has  appar- 
ently a  complete  loss  of  yellow.  The  green-yellow-red  blotched  and  the 
green-yellow  spotted-red  blotched  patterns  are  gradations  between  these 
extremes. 

Color  pattern  yellow-green-red  blotched  (fig.  6) :  This  is  a  pattern 
of  green,  yellow,  and  red  as  in  type  green-yellow-red  blotched,  but  the 
relative  positions  of  the  green  and  the  j^ellow  are  reversed.  The 
3^ellow  is  in  the  central  portion  of  the  leaf. 

Color  pattern  green-yelloir-solid  red  (fig.  8) :  Both  surfaces  of  the 
leaf  are  a  solid  red.  Through  the  center  of  the  leaf  the  color  is 
violet  carmine,  but  the  marginal  zone  underlaid  by  j^ellow  is  nopal  red. 
At  the  base  of  the  leaves  a  greenish  tint  prevails  and  at  the  extreme 
edge  of  the  margin  a  fine  line  of  yellow  is  visible.  On  the  under  sur- 
face the  red  seems  slightly  less  intense  and  does  not  cover  the  larger 
veins,  which  stand  out  prominently^  on  this  surface.  This  pattern 
differs  from  green-yellow-red  blotched  in  having  the  entire  epidermis 
solid  red  instead  of  blotched.  Frequently,  however,  a  few  isolated 
areas  are  free  of  epidermal  red  and  the  underlying  green  or  yellow 
shows  clearly. 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  19 

Color  pattern  green-yellow  spotted-solid  red:  This  pattern  has  the 
solid  red  as  in  the  preceding  type,  but  the  conditions  of  yellow  and 
green  are  as  in  type  green-yellow  spotted-red  blotched. 

Color  pattern  green-solid  red  (fig.  9):  This  pattern  has  the  entire 
leaf  above  and  below  of  a  uniform  violet  carmine.  It  differs  from  type 
green-yellow-solid  red  in  the  absence  of  any  underlying  yellow,  and  from 
type  green-red  blotched  in  having  the  epidermis  completely  red  instead 
of  red  in  blotches.  The  pattern  is  dull  and  dark,  with  a  somewhat 
metallic  luster,  in  marked  contrast  to  the  various  patterns  with  yellow. 

Color  pattern  green-yellow-solid  red  upper  center  (figs.  10  and  lOo) : 
This  is  a  brightly  colored  and  attractive  pattern  with  a  rather  com- 
plicated arrangement  of  colors.  The  subepidermal  colors  of  green 
center  and  yellow  border  are  similar  to  the  arrangement  in  the  types 
green-yelloiv-red  blotched  and  green-yellow-solid  red.  The  epidermal 
red  is,  however,  almost  entirely  confined  to  the  upper  surface;  over 
the  central  green  it  gives  a  greenish  violet  carmine  cast;  over  the  border- 
ing yellow  it  forms  a  band  of  nopal  red.  At  the  extreme  margin  it  is 
absent,  giving  a  narrow  but  irregular  band  of  pure  yellow.  On  the 
under  surface  there  are  only  occasional  small  blotches  of  red.  About 
the  border  the  red  of  the  upper  epidermis  shows  through  the  yellow, 
giving  pale  pinkish  tints,  as  shown  in  figure  10a.  This  type  was 
derived  from  pattern  green-yellow-solid  red,  with  its  complete  covering 
of  epidermal  red,  by  the  loss  of  red  on  the  under  surface  and  about 
the  extreme  margin  of  the  upper  surface. 

Color  pattern  yelloiv-green-solid  red  (fig.  11):  This  pattern  w^as 
derived  from  type  green-yellow-solid  red  by  a  reversal  in  the  relative 
position  of  the  underlying  green  and  yellow,  the  change  being  the  same 
that  gave  type  yellow-green-red  blotched  from  pattern  green-yellow-red 
blotched. 

Color  pattern  green-yellow  (fig.  12) :  This  is  a  bright  pattern  of  green 
center  and  yellow  border  with  no  expression  of  epidermal  red.  Some  few 
internal  or  vascular  strands  of  red  may  be  seen.  The  pattern  difi'ers 
from  that  of  type  green-yellow-red  blotched  in  having  no  epidermal  red. 

Color  pattern  green  (fig.  13) :  A  pattern  of  pure  spinach  green  with 
no  yellow  and  no  epidermal  red,  but  with  a  few  streaks  of  red  in  the 
vascular  strands  or  in  the  mesophyl.  This  pattern  difi'ers  from  the 
parental  pattern  green-yellow-red  blotched  in  the  loss  of  both  yellow^  and 
epidermal  red.  The  green-yellow  spotted  pattern  (type  13  a,  not  illus- 
trated) differs  only  in  having  yellow  spots. 

Color  pattern  yellow-green  (fig.  14) :  This  type  has  a  green  border 
and  a  yellow  center,  with  no  epidermal  red.  It  differs  from  the 
green-yellow  pattern  in  the  reversed  position  of  the  two  color  elements 
and  from  pattern  yellow-green-red  blotched  in  having  no  epidermal  red. 
As  in  the  case  of  the  green-yellow  type,  the  pattern  is  bright  and 
attractive. 


20  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

Color  pattern  green-solid  red  upper  center  (fig.  15):  A  type  that 
differs  from  type  green-yellow-solid  red  upper  center  in  having  no  yellow 
and  hence  is  apparently  bicolored  on  the  upper  surface.  The  center 
of  the  leaf  is  violet  carmine  and  the  marginal  zone  is  pure  green. 

Color  pattern  yellow-solid  red:  This  tj^pe  has  almost  uniform  nopal 
red  color  on  both  surfaces.  It  differs  from  type  green-yellow-solid  red 
in  not  possessing  a  dark  red  center  and  from  type  yellow-red  blotched 
in  having  the  epidermis  completely  red.  In  both  patterns  the  green 
underlying  a  solid  red  epidermis  is  almost  entirely  absent. 

Laciniate  leaf  shape  (fig.  7) :  In  marked  contrast  to  the  type  of  entire 
leaf  illustrated  in  the  figures  showing  the  various  color  patterns  is 
the  deeply  and  irregularly  cut  and  lobed  types  of  leaf  shape,  the  appear- 
ance and  behavior  of  which  will  be  specially  discussed  later. 

All  of  these  color  patterns  arose  as  sudden  spontaneous  bud-varia- 
tions, with  the  single  exception  of  the  type  green-solid  red  upper  center, 
which  is  a  pattern  into  which  plants  with  the  green-yellow-solid  red 
upper  center  pattern  fluctuated.  Throughout  this  paper,  as  above 
noted,  the  term  "bud  variation"  is,  in  all  cases  not  otherwise  quahfied, 
apphed  only  to  a  marked  change  that  appeared  suddenly  and  com- 
pletely for  a  part  of  a  plant,  and  which  was  fully  in  evidence  in  the 
leaves  involved  when  they  first  unrolled.  Gradual  fluctuations  also 
gave  in  numerous  cases  types  green-yellow  spotted-red  blotched,  green- 
yelloiv  spotted,  and  green-red  blotched.  That  is,  these  types  appeared 
both  by  sudden  and  by  gradual  variations. 

CONSTANCY  OF  THE  VARIOUS  PATTERNS. 

To  test  the  constancy  of  the  t>T)es,  the  original  as  well  as  those 
derived  from  it  by  bud  variations,  successive  generations  of  plants 
were  grown  from  pedigreed  cuttings.  This  tested  the  vegetative  con- 
stancy of  the  pattern  itself  and  enables  one  to  make  comparisons  when 
the  same  pattern  was  derived  from  different  lines. 

The  series  of  plants  considered  under  any  type  pattern  are  in  large 
measure  a  selected  stock.  When  cuttings  were  made  for  the  perpetua- 
tion of  the  pattern  in  a  new  generation,  they  w^ere  made  from  the  plants 
most  typical  and  constant  for  the  pattern  concerned.  \\Tien  a  bud 
variation  appeared,  if  the  conditions  were  favorable,  the  parts  posses- 
sing it  were  allowed  to  develop  until  there  were  several  branches  from 
which  cuttings  could  be  taken  simultaneously.  In  such  cases  the 
selection  of  branches  for  the  new  type  was  a  simple  matter,  as  it 
depended  on  the  taking  of  branches  sharply  distinct  from  the  main 
part  of  the  plant,  which  in  most  cases  were  as  different  as  is  shown  in 
figures  21  and  24.  When  further  cuttings  were  made  for  a  new  genera- 
tion to  perpetuate  the  type  they  were  made  from  plants  most  uniform 
and  constant  (determined  from  the  records)  for  the  pattern  in  question. 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  21 

Usually  but  three  cuttings  were  made  from  a  plant,  and  these  were 
taken  from  branches  most  uniform  and  clearly  conforming  to  the  type. 

It  has  already  been  noted  and  it  will  be  very  evident  in  the  following 
pages  that  some  plants  showed  fluctuating  variations  giving  irregular 
or  mixed  patterns,  or  the  pattern  gradually  fluctuated  between  two 
types  or  changed  from  one  type  to  another.  Except  in  two  cases 
no  attempt  was  made  to  secure  new  types  by  such  fluctuating  varia- 
tions. These  cases  (clone  14  of  table  2  and  clone  13  of  table  3)  will 
be  especially  discussed  later. 

In  numerous  cases  cuttings  were  made  to  give  two  types  of  patterns 
in  the  same  plant.  The  constancy  of  the  patterns  could  in  these  cases 
be  studied  with  the  two  parts  growing  from  the  same  root  system  and 
submitted  to  the  same  environmental  factors. 

Plants  with  yellow-red  blotched  pattern  (fig.  1). — Seven  cuttings  pure 
for  this  pattern  were  made  in  the  autumn  of  1913.  Six  died  soon  after 
they  were  placed  in  the  rooting-bench.  The  other  lived  and  was  grown 
until  the  autumn  of  1914.  This  plant  was  somewhat  greener  during 
the  winter,  but  at  all  times  was  decidedly  more  yellow  than  any  plant 
of  any  other  type.  It  was,  also,  smaller  and  less  vigorous  in  its  growth. 
Eight  plants  were  grown  as  chimeras  with  one  branch  of  yellow-red 
UotchedpSitteTn  and  one  branch  of  a  pattern  with  yellow-green-red  blotched. 
On  all  these  the  branches  of  the  part  with  pattern  yellow-red  blotched 
remained  quite  constant  throughout  the  year  and  were  at  all  times  in 
marked  contrast  to  the  pattern  of  the  other  part.  Two  chimeras  grown 
only  during  the  summer  of  1914  were  likewise  quite  constant.  While 
it  is  very  difficult  to  obtain  plants  with  this  pattern  from  cuttings,  the 
type  remains  quite  constant  when  grown  in  chimeral  association  with 
branches  having  green  tissue.  On  account  of  the  difficulties  of  propa- 
gation this  type  has  not  been  rigorously  tested.  The  few  plants  grown 
gave  no  marked  variations  either  as  bud  variations  or  as  fluctuations. 

Plants  with  green-yellow-red  blotched  pattern  (fig.  2). — This  is  the 
pattern  originally  possessed  by  the  two  parent  plants,  Nos.  1  and  3. 
A  total  of  337  plants  were  grown  from  cuttings  of  this  type.  The  data 
given  in  table  2  are  summarized  in  four  main  clones.  Plants  of  clone  1 1 
all  descended  from  branch  1  on  plant  1.  Plants  of  clone  12  were 
descended  from  branch  2  on  plant  1.  Plants  of  clone  3  were  derived 
from  the  branches  of  plant  3  that  were  uniform  for  this  pattern.  The 
original  branches  from  which  the  first  cuttings  were  obtained  were 
uniform  for  the  green-yelloiu-red  blotched  pattern  and  all  plants  used  as 
parent  stock  for  later  generations  were  selected  as  typical  and  most 
constant  for  the  pattern.  The  entire  six  generations  constituted  a 
series  of  plants  derived  by  continued  selection. 

The  45  plants  of  this  pattern  in  clone  14  are  especially  interesting, 
as  they  constitute  a  test  for  this  pattern  when  derived  by  a  gradual 
fluctuation.    As  already  noted,  branch  4  of  plant  1  possessed  a  decidedly 


22 


ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 


green  pattern  with  yellow  blotches  designated  as  green-yellow  spotted-red 
blotched.  In  the  third  generation  of  plants  grown  from  this  branch 
8  plants  gradually  fluctuated  during  winter  until  they  were  uniformly 
green-yellow-red  blotched.  Such  an  increase  of  yellow  during  winter  was 
unusual  and  cuttings  were  made  to  test  the  constancy  of  the  type  thus 
derived.  The  data  for  the  45  plants  grown  in  three  generations  show 
that  two-thirds  of  the  plants  were  quite  constant  for  the  derived  type. 
This  proportion  compares  very  favorably  v/ith  that  of  the  clones  11,  12, 
and  3,  which  were  from  the  start  selected  from  plants  most  typical  for 
the  type. 

Table  2. — Summary  of  plants  irith  green-yellow-red  blotched  ■pattern. 


Clone 
11. 


Clone 
12. 


Clone 
14. 


Clone 
3. 


Total. 


Number  of  plants 

Number  constant 

Number  fluctuating  in  green  and  yellow 
Fluctuations  to  laciniate  leaf-shape .... 

Plants  giving  bud  variations 

Bud  variations: 

Yellow-red  blotched 

Green-yellow  spotted-red  blotched . 

Green-red  blotched 

Yellow-green-red  blotched 

Green-yellow-solid  rod 

Green-j'ellow 

Green-yellow  spotted 


151 

102 

29 

22 


4 

13 

3 


103 

76 

22 

1 

12 

1 
2 

7 

1 
1 


45 
21 
15 

10 

3 

14 
1 
2 

1 
1 

2-? 


38 

19 

11 

1 

9 


337 
218 

77 
2 

53 

7 
8 
24 
6 
4 
8 
o 


'All  were  cases  of  fluctuation  confined  to  about  half  of  a  plant. 

^Loss  of  epidermal  red  on  two  plants  fluctuating  from  yellow  to  yellow  spotted. 

Of  the  total  number  of  plants  with  pattern  green-yelloiv-red  blotched 
there  were  218  that  were  at  all  times  fairh^  constant  and  true  to  the 
type.  They  were  all  somewhat  fluctuating  in  respect  to  the  relative 
amounts  of  green  and  yellow,  but  were  all  constant  in  possessing  at  all 
times  a  yellow  border. 

In  figure  2  there  is  shown  a  leaf  with  the  average  development  of 
the  yellow  border,  although  in  this  leaf  the  pattern  is  somewhat  irregular. 
Figures  10,  12,  19,  20,  and  26  show  leaves  classed  as  yellow-bordered; 
figure  20,  however,  shows  fluctuation  toward  tyipe yellow-red  blotched,  and 
26  shows  fluctuation  toward  a  green-yellow  spotted-red  blotched  pattern. 
Figures  17  and  27  are  from  leaves  classed  as  having  irregular  patterns. 

The  77  cases  classed  as  fluctuations  include :  (1)  56  cases  of  decided 
increase  of  green  during  winter,  followed  by  increase  of  yellow  in 
summer,  giving  in  most  cases  return  to  the  type  of  the  cutting;  two  of 
these  also  gave  fluctuations  to  laciniate-leaf  shape;  (2)  9  cases  of 
increase  of  yellow  during  summer  (grown  onlj^  during  a  summer  from 
cuttings  taken  from  plants  that  were  greener  during  the  preceding 
winter) ;  (3)  7  cases  of  fluctuations  that  were  not  uniform  on  a  plant, 
but  gave  leaves  of  the  same  age  with  different  patterns  so  mixed  that 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  23 

no  sectorial  distribution  could  be  traced;  and  (4)  5  plants  with  green 
and  yellow  distributed  irregularly. 

Aside  from  the  fluctuations  in  relative  amounts  of  green  and  yellow, 
there  was  also  much  fluctuation  in  number  and  size  of  the  l)lotches  of 
epidermal  red.  On  some  of  the  plants  there  was  rather  gradual  increase 
or  decrease  both  in  number  and  size  of  these  blotches,  giving  such 
differences  in  respect  to  red  as  are  shown  in  figures  5,  17,  26,  and  28. 
Such  plants  were,  however,  still  considered  as  blotched  in  the  sum- 
maries. Selection  of  typical  red  blotched  epidermis  for  various  types 
has  been  directed  to  plants  having  the  epidermal  blotching  as  in  figures 
2,  5,  and  6,  rather  than  as  in  figures  23  and  28. 

53  plants  produced  bud  variations  giving  loss  of  green  7  times,  loss 
of  the  yellow  bordering-band  8  times,  complete  loss  of  yellow  24  times, 
reversal  of  the  relative  positions  of  the  green  and  yellow  6  times  (4  cases 
appeared  in  half  of  a  leaf  only,  as  in  fig.  25),  increase  of  red  to  com- 
plete epidermal  red  4  times,  complete  loss  of  epidermal  red  10  times. 
In  clone  14,  4  plants  gradually  developed  a  green-yelloiv  spotted-red 
blotched  pattern  in  part  of  the  branches.  On  the  basis  of  my  descrip- 
tions the  change  was  a  fluctuating  variation  affecting  only  a  few 
branches  of  a  plant.  These  4  cases  are  not  included  in  the  summaries 
of  bud  variations. 

In  their  extent  the  bud  variations  gave  extremes  in  development  of 
yellow,  of  green,  and  of  epidermal  red.  There  were  cases  of  nearly 
pure  yellow  and  of  absolutely  pure  green ;  there  were  cases  of  solid  red 
epidermis  and  others  with  no  red  epidermis. 

Furthermore,  the  changes  in  green  and  yellow  or  in  epidermal  red 
occurred  entirely  independentl}^  of  each  other.  In  general,  the  different 
types  of  bud  variations  were  quite  uniformly  distributed  in  the  various 
clones. 

The  type  green-yellow  spotted  was  produced  on  2  plants  by  a  fluctua- 
tional  increase  of  green  after  the  loss  of  epidermal  red  had  occurred. 

In  table  2,  as  in  other  tables,  when  the  totals  given  for  constant, 
fluctuating,  and  sporting  plants  exceed  the  number  of  plants  grown, 
it  shows  that  a  certain  number  of  the  fluctuating  plants  produced  also 
sharp,  clear-cut  bud  variations.  Also,  when  the  total  of  cases  of  bud 
variations  exceeds  the  plants  giving  them,  certain  plants  produced 
more  than  one  bud  variation. 

For  the  purpose  of  estabhshing  an  index  of  the  frequency  of  bud 
variation  we  may  take  the  ratio  of  bud  variations  to  the  estimated 
number  of  buds  developed.  Each  plant  produced  an  average  of  at 
least  200  branches  which  made  sufficient  growth  to  reveal  the  pattern 
of  the  leaves.  On  this  basis  the  index  of  total  bud  variation  for  this 
group  was  about  1  to  1,110.  The  ratio  of  constant  plants  to  fluctu- 
ating plants  was  almost  exactly  3  to  1,  not  counting  the  plants  with  bud 
variations  many  of  which  were  otherwise  constant. 


24 


ESTABLISHMENT    OF   VARIETIES   IN    COLEUS 


Plants  with  color  pattern  green-yellow  spotted-red  blotched  and  with 
uniformly  entire  leaves. — The  plants  grouped  in  this  class  (figs.  4  and 
23)  present  perhaps  greater  diversity  than  those  of  any  other  type, 
embracing  (1)  plants  with  considerable  yellow  in  scattered  areas  in  all 
leaves,  (2)  plants  with  only  shght  amounts  of  yellow  in  scattered  areas 
in  nearly  all  leaves,  and  (3)  plants  with  only  a  few  leaves  possessing  yel- 
low spots.  Between  the  extremes  there  was  every  degree  of  variation 
and  often  all  degrees  would  be  seen  at  one  time  among  the  leaves  of  a 
single  plant. 

It  is  difficult  in  such  plants  to  determine  what  constitutes  a  variation 
either  as  a  fluctuating  or  a  bud  variation  when  it  involves  green  and 
yellow.  The  cases  given  in  table  3  are  those  in  which  an  entire  branch 
or  a  sector  of  a  branch  showed  leaves  that  were  uniform  for  a  new 
pattern.  Plants  having  irregular  mixtures  of  leaves  of  equally  different 
patterns  were  common.     Such  cases  are  of  special  interest,  as  are  the 

Table  3. — Plants  irith  entire  leanes  and  pattern  green-yellow  spotted-red  blotched  {fig.  4)- 


Clone 
11. 


Total  number  of  plants 

Number  constant 

Fluctuations  for  green  and  yellow 

To  type  green-yellow-red  blotched  .  .  . 

To  mixed  patterns 

To  laciniate  leaf 

Total  number  plants  gi\nng  bud  variations . 
Bud  variations: 

To  yellow-red  blotched 

To  green-yellow-red  blotched 

To  green-red  blotched 

To  green-yellow  spotted-solid  red .  .  .  . 

To  green-yellow  spotted 


Clone      Clone 
12.  13.1 


16 
6 
2 

2 
4 
2 


79 
01 
16 


Clone 
14. 


89 
50 
21 
8 
4 
3 
3 

1 
1 
2 


Clone 
3. 


22 
9 

8 

4 
1 


Total. 


198 

126 

47 

8 
13 

7 
8 

1 
3 

2 
2 
2 


^  Pattern  derived  by  fluctuating  variation. 

fluctuating  variations  in  a  seed  progen5\  They  possess  many  simi- 
larities to  cases  of  size  inheritance  described  b}^  Goodspeed  (1912)  and 
raise  the  question  as  to  whether  color  heredity  is  not  also  quantitatively 
rather  than  qualitatively  inherited. 

As  has  already  been  noted,  the  parent  plant  (No.  1)  had  one  branch 
(No.  14)  with  leaves  green-yellow  spotted-red  blotched.  All  the  89  plants 
of  clone  14  descended  from  this  branch  through  6  generations  of  selec- 
tion. The  plants  of  this  pattern  here  given  with  clones  11,  12,  and  3 
were  obtained  from  cases  of  bud  variation  from  the  type  green-yellow- 
red  blotched  (see  table  2).  The  79  plants  of  clone  13,  however,  were 
derived  from  5  plants  that  gave  a  fluctuating  change  from  green-red 
blotched  to  green-yellow  spotted-red  blotched.  This  was  a  frequent 
fluctuation  from  the  green  plants  especially  of  clone  13,  as  shown  in 
table  5,  and  the  yellow-spotted  condition  thus  obtained  was  tested  in 
four  generations,  comprising  a  total  of  79  plants. 


BY   THE    SELECTION    OF    SOMATIC   VARIATIONS.  25 

As  a  whole,  there  was  a  rather  large  proportion  of  the  plants  that 
remained  within  the  type  as  classified,  although  there  was  hardly  a 
plant  grown  during  the  winter  that  did  not  become  somewhat  greener 
during  that  period.  68  plants  fluctuated  in  a  marked  degree;  47  of 
them  were  almost  entirely  green  during  the  winter,  but  were  again 
quite  uniformly  yellow  spotted  in  summer,  although  some  of  these 
remained  much  greener  in  summer.  None  of  the  latter,  however,  could 
be  considered  as  of  the  pattern  green-red  blotched. 

13  plants  fluctuated  irregularly,  giving  mixed  patterns,  mostly  of 
green-yellow-red  blotched,  green-yellow  spotted-red  blotched,  and  green-red 
blotched,  all  more  or  less  intermingled  among  the  various  branches  and 
on  the  same  branch.  These  were  not  used  as  parent  plants,  but  doubt- 
less by  selection  it  would  be  possible  to  obtain  a  marked  degree  of 
constancy  for  the  irregular  and  mixed  patterns,  with,  also,  production 
of  plants  that  would  be  uniform  for  various  types. 

The  most  uniform  and  marked  of  the  fluctuations  was  the  case 
of  8  plants  of  clone  14  which  gradually  became  more  yellow  during 
the  winter  of  1912-13,  until  they  were  quite  typical  green-yellow-red 
blotched;  5  of  these  were  used  as  parents  of  the  plants  of  clone  14  already 
reported  with  table  3.  The  change  in  pattern  arose  as  a  gradual 
increase  of  yellow  from  various  degrees  of  a  yellow  spotted  condition  to 
a  well-defined  yellow  bordered  pattern  that  was  quite  uniform  for  the 
entire  plant,  and  which  when  tested  in  progeny  was  subsequently  quite 
as  constant  as  cases  which  arose  by  sudden  variations.  An  analysis  of 
the  pedigrees  of  these  8  cases  shows  that  all  of  these  descended  from 
only  3  of  the  7  plants  grown  from  cuttings  of  the  original  branch  14. 
This  phenomenon  of  the  appearance  of  the  same  variation  in  different 
plants  that  were  derived  from  the  same  more  remote  ancestor  is  common 
and  constitutes  what  we  may  call  duplicate-reversions  or  variations. 

Besides  the  fluctuating  variations  in  regard  to  green  and  yellow, 
there  were  numerous  cases  of  fluctuation  in  the  red-blotched  condition 
both  of  the  epidermis  and  of  the  subepidermal  tissues,  giving  extremes 
of  very  finely  red-blotched  or  coarsely  blotched.  No  selections  were 
made  to  secure  types  of  the  red-blotched  condition.  None  of  the 
plants  fluctuated  to  a  no-blotched  or  to  a  soUd-red  pattern. 

The  behavior  of  the  79  plants  of  this  pattern  in  clone  13  is  especially 
interesting.  They  constitute  a  test  for  this  pattern  obtained  by  the 
selection  of  gradual  and  accumulated  fluctuation.  The  progeny  of 
5  plants  grown  in  3  generations,  subjected  to  the  same  sort  of  selection 
as  the  other  clones,  showed  the  highest  degree  of  constancy  obtained  in 
the  clones  of  this  pattern. 

Seven  plants  gave,  during  the  winter,  a  marked  increase  of  green,  ac- 
companied by  the  production  of  cut  and  laciniate  leaves  (fig.  7),  the 
appearance  and  constancy  of  which  are  quite  fully  discussed  later. 


26  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

There  were  few  cases  of  sudden,  clear-cut  bud  variation  in  this 
group.  One  was  a  very  decided  and  ahnost  complete  development  of 
yellow  to  yellow-red  blotched;  3  were  sectorial  variations  to  green-yellow- 
red  blotched;  2  cases  gave  loss  of  yellow  to  green-red  blotched;  2  involved 
gain  of  epidermal  red  to  solid  red,  and  in  two  cases  there  was  loss  of 
epidermal  red.  All  these  cases  were  sectorial  for  a  plant  or  in  some 
cases  for  a  single  branch.  The  ratio  giving  the  frequency  of  bud  varia- 
tions involving  color  in  this  group  is  1  to  3,960. 

Plants  with  laciniate  leaves. — Until  the  winter  of  1912-13,  all  the 
plants  in  the  cultures  had  been  constant  and  very  uniform  for  leaf- 
shape,  showing  no  greater  variation  in  this  respect  that  is  seen  in  fig- 
ures 2,  6,  and  9.  During  that  Mdnter  it  was  noted  in  11  cases  that  as 
new  leaves  developed  they  were  more  and  more  deeply  cut  and  lobed 
until  in  Januar}^  and  February  the  uppermost  leaves  were  in  extreme 
cases  much  divided  and  deeply  laciniate,  as  shown  in  figure  7.  The 
plants  appeared  like  the  middle  plant  in  plate  4;  3  of  these  plants 
were  from  cuttings  of  branches  that  were  pure-green  bud  variations; 
the  others  were  plants  which  had  a  somewhat  fluctuating  green-yellow 
spotted  pattern.  Nine  of  these  plants  were  grown  during  the  following 
summer,  when  it  was  noted  that  without  exception  the  new  leaves 
gradually  became  more  entire  until  by  late  summer  all  the  leaves  then 
hanging  to  the  plants  were  entire.  At  the  same  time  5  of  the  plants 
became  decidedly  more  yellow,  even  becoming  quite  uniform  for  type 
green-yellow-red  blotched.  Cuttings  were  made  in  April  from  each  of 
these  plants  and  from  these  16  plants  were  grown  during  the  summer 
of  1913.  All  of  these  fluctuated  to  entire  leaves,  and  in  regard  to  color 
gave  plants  some  of  which  were  uniform  for  type  green-yellow-red 
blotched,  others  for  tj^pe  green-yellow  spotted-red  blotched,  while  the 
foliage  of  others  showed  mixtures  of  these  patterns  with  also  pattern 
green-red  blotched. 

The  further  generations  in  subclone  12  exhibited,  as  shown  in  table 
4,  the  same  periodicity  in  change  of  leaf-form,  except  that  in  late  sum- 
mer of  1914  a  rather  large  number  of  the  plants  showed  new  leaves 
that  were  laciniate.  One  plant  of  this  subclone  was  grown  in  the  winter 
of  1913-14  from  a  pure-green  bud-sport  that  appeared  during  the 
previous  summer.  This  plant  remained  constant  for  entire  leaves  of 
pattern  green-red  blotched  during  the  winter,  but  as  it  died  early  in  the 
summer  no  further  progeny  were  grown.  Further  generations  of  the 
plants  with  laciniate  leaves  in  subclone  14  were  not  grown. 

The  laciniate  leaf-form  appeared  anew  in  the  winter  of  1913-14  in 
clone  3  in  two  instances.  One  was  a  plant  whose  line  of  descent 
showed  bud  variation  from  type  green-yellow-red  blotched  to  type  green- 
yellow  spotted-red  blotched;  that  of  the  other  showed  bud  variation  from 
green-yelloio-red  blotched  to  green-yellow  solid  red,  and  then  from  this  to 
green-yellow-solid  red  upper  center.     The  latter  was  the  only  one  of 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS. 


27 


8  plants  grown  from  cuttings  of  a  single  plant  to  exhibit  the  fluctuation 
to  laciniate  leaves. 

It  is  to  be  noted  that  this  character  of  laciniate  leaf-shape  has  not 
appeared  thus  far  in  any  of  the  plants  grown  in  clones  11  and  13.  It 
has  appeared  as  a  fluctuating  character  that  develops  most  strongly  in 
winter.  With  one  exception  all  the  plants  grown  to  test  the  reappear- 
ance of  the  character  have  exhibited  it.  This  plant  was  grown  from 
a  bud  variation  giving  a  single  branch  of  green-red  blotched  on  a  plant 
otherwise  uniform  for  pattern  green-yellow-red  blotched  at  the  time  the 
cutting  was  made. 

During  the  time  these  plants  exhibited  the  laciniate  character  most 
strongly,  there  were  growing  among  them  numerous  plants  of  other 
clones  of  various  patterns,  especiall}'-  of  green-yelloir-red  blotched  and 
green-red  blotched,  all  submitted  to  the  same  conditions  of  Ught,  tem- 

Table  4. — Summary  of  plants  with  laciniate  leaves. 


Total  number  of  plants 

Much  greener  in  winter .... 
Very  laciniate  in  ^\dnter.  .  .  . 

Entire  in  winter 

In  summer  as  tjije  2 

as  type  4 

as  type  1 

mixed  patterns . . 

uniformly  entire. 

slightly  laciniate . 


Plants  giving  bud  variations 

Type  green-red  blotched  and  entire . 

green-yellow  spotted 

green-yellow-red  blotched 


Clone  12. 


1912- 
1913, 


6 
f) 
G 
0 
5 
0 
0 
1 
6 
0 

1 
1 


1913. 


10 


4 
1 
0 
5 
10 
0 


1913- 
1914. 


32 

32 

31 

1 

4 

.5 

1 

22 

5 

27 

1 

1 


1914. 


1 
3 
0 
3 

0 

7 


Clone  14. 


1912- 
1913. 


1913. 


3 
3 
3 
0 
0 
0 
0 
3 
3 
0 


0 
6 
0 
0 
6 
0 


Clone  3. 


1913- 
1914. 


1914. 


2 
2 
2 
0 
0 
0 
0 
2 
1 
1 


0 
1 

1 
0 
0 
2 


perature  and  soil,  yet  not  in  the  least  degree  exhibiting  the  fluctuation 
to  laciniate  leaves.  This  is  well  shown  in  plate  4,  which  gives  a  photo- 
graph of  3  plants  of  the  same  clone  (12),  all  grown  under  the  same 
conditions.  The  plant  to  the  right  (No.  125111)  had  entire  leaves 
and  a  green-yellow-red  blotched  pattern;  the  one  to  the  left  (No.  1251412) 
was  of  a  pure  green-red  blotched  pattern;  the  one  in  the  middle  (No. 
123153)  shows  the  transition  from  entire  leaves  to  deeply  laciniate 
leaves  as  it  occurs  during  the  winter. 

The  late  summer  of  1914  was  exceedingly  dry.  In  July  there  had 
been  5.36  inches  of  rainfall,  during  which  time  the  plants  made  an 
unusually  vigorous  growth.  From  August  12  until  October  16  there  was 
but  1.26  inches  of  rain.     During  the  dry  warm  period  of  September,  the 


28 


ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 


new  leaves  on  very  many  of  the  plants  of  this  series  were  strongly 
laciniate.  Cuttings  were  made  from  these  laciniate-leaved  branches. 
The  new  leaves  that  developed  on  these  young  plants  during  November 
were  entire;  hence  it  would  seem  that  the  laciniate  character  in  these 
particular  clones  of  Coleus  is  in  some  degree  associated  with  decreased 
vigor.  When  most  favorable  conditions  for  growth  prevail,  or  when 
rapid  growth  is  brought  about  in  cuttings,  the  leaves  become  entire. 

It  is,  however,  clear  that  the  first  appearance  of  the  laciniate  char- 
acter was  confined  to  a  few  plants  and  that  once  it  originated  it 
reappeared  with  marked  constancy  in  the  vegetative  progeny. 

During  the  winter  of  1914-15  the  laciniate  character  appeared  in 
the  manner  of  a  bud  variation.  A  large  plant  that  had  been  grown  out 
of  doors  during  the  summer  was  in  September  severely  pruned  and 
placed  in  a  pot  for  further  development  in  the  greenhouse.  It  was 
intended  to  use  the  plant  for  stock  in  general  border  planting  and  the 
plant  label  was  not  preserved.  From  the  records  of  the  pattern  and 
generation  it  is  clear  that  the  plant  itself  and  all  the  plants  in  its  line 
of  descent  possessed  only  entire  leaves  and  the  plant  belonged  to  the 
main  clone  1.     This  plant  was  given  the  number  9. 

In  the  course  of  3  months  numerous  new 
branches  developed  on  the  10  pairs  of  main 
lateral  branches  to  which  the  plant  had 
been  pruned.  It  was  noticed  that  of  the 
20  main  branches,  3  bore  branches  with 
laciniate  leaves.  The  positions  of  these  are 
indicated  by  numbers  1,  2,  and  3  of 
diagram  2. 

All  the  branches  on  all  other  of  the  main 
branches  bore  only  entire  leaves.  All  the 
branches  arising  from  2  bore  only  laciniate 
leaves,  but  the  branches  with  entire  and 
with  laciniate  leaves  were  sectorially  dis- 
tributed on  the  branches  1  and  3.  The 
contrast  was  most  marked,  especially  when 
opposite  branches  were  different,  one  hav- 
ing laciniate  and  the  other  entire  leaves.  The  sectorial  differences 
appeared  in  some  of  the  secondary  branches  and  carried  the  two  types 
into  parts  of  individual  leaves. 

The  most  striking  behavior  of  this  series  of  plants  summarized  in 
table  4  is  the  wide  fluctuations  in  the  leaf-shape  and  in  the  amount  of 
yellow  and  green,  the  marked  correlation  of  decrease  of  yellow  with 
decrease  of  leaf  area,  and  the  rather  pronounced  periodicity  of  these 
fluctuations.  These  fluctuations  are  so  general  and  rhythmic  that 
they  can  almost  be  considered  constant.  Of  bud  variations  there  were 
but  3  cases,  giving  a  ratio  of  about  1  to  2,530,  which,  however,  shows 


10 

10' 

9 

9' 

8 

8' 

7 

7' 

6 

6' 

5 

5' 

4 

4' 

3 

3' 

2 

2' 

1 

1' 

Diagram  2.  —  Showing  position 
of  the  branches  on  plant 
No.  9. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS. 


29 


greater  frequency  than  that  of  the  pattern  green-yellow  spotted-red 
blotched.  Of  the  bud  variations  one  was  a  loss  of  yellow,  one  gave 
increase  of  yellow  to  type  green-yelloiv-red  blotched  and  one  was  a  loss 
of  red. 

Plants  ivith  green-red  blotched  -pattern. — The  apparently  complete  loss 
of  yellow,  giving  only  green  subepidermal  tissues,  was  a  frequent  bud 
variation  in  plants  having  green-yellow,  yellow-green,  or  green-yellow 
spotted  patterns,  regardless  of  the  degree  of  red  in  the  epidermis  (fig.  5). 
The  condition  of  pure  green  also  developed  as  a  fluctuation  on  plants 
of  these  same  types.  In  cases  the  fluctuation  was  quite  general  for  the 
entire  plant,  while  in  others  it  occurred  irregularly,  giving  plants  with 
mixed  patterns. 

The  90  plants  included  in  this  summary  are,  however,  selected  stock, 
all  descended  from  cases  of  bud  variation  similar  to  that  of  figure  21, 
in  which  the  part  concerned  showed  no  trace  of  yellow  in  any  leaves. 

Table  5. — Summary  of  plants  unth  green-red  blotched  pattern  (fig.  5). 


Clone  j  Clone 
11.  12. 


Clone  I  Clone 
13.      1      14. 


Clone 
.3. 


Total. 


Number  of  plants 

Plants  constant 

Fluctuations: 

Green-yellow  spotted 

Mixed  patterns 

Laciniate  leaf-shape 

Plants  with  bud  variations 

Bud  variations: 

Green-yellow  spotted-red  blotched .  . 

Spontaneous  yellow 

Green-solid  red 

Green 


13 
5 


1 
6 


18 
5 

7 
1 
3 
5 

1 
1 
2 
1 


51 
17 

31 
3 


6 
3 


2 
1 


90 
31 

41 
4 
3 

15 

2 
9 
2 
3 


Selections  for  further  generations  were  made  from  plants  that  had 
remained  uniformly  pure  green.  The  type  was  maintained  bj^  selection 
quite  as  it  is  practiced  in  a  herd  of  dairy  cattle.  In  the  case  of  clone  13, 
6  generations  were  grown,  all  descended  from  the  pure-green  branch  of 
the  parent  plant,  No.  1. 

Of  the  total  number  of  plants  in  this  group,  31  remained  pure  green, 
showing  no  trace  of  yellow  by  either  fluctuation  or  bud  variation. 
In  addition,  14  of  the  15  plants  with  bud  variations  were  otherwise 
constant  for  the  pure-green  condition.  All  of  these  plants  were  grown 
during  an  entire  summer.  41  plants  developed  varying  amounts  of 
yellow  in  scattered  areas,  making  a  pattern  classed  as  green-yellow 
spotted-red  blotched.  In  11  of  these  the  yelloio  spotted  condition  was 
quite  uniform  and  typical  and  from  these  were  selected  parents  for 
the  plants  of  subclone  13  given  in  the  table  3.  Three  plants  also  gave 
fluctuations  in  leaf-shape  to  the  laciniate  type  and  their  progeny  are 
included  in  the  summary  of  table  4.     Four  plants  gave  decidedly 


30 


ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 


mixed  and  irregular  patterns  like  those  that  appeared  from  patterns 
treated  in  tables  3  and  4. 

There  were  16  cases  of  bud  variation;  5  were  concerned  with  the  red 
epidermis,  2  giving  solid  red,  and  3  giving  no  red.  There  were  2  cases 
of  vspontaneous  appearance  of  yellow,  giving  branches  sectorial  for  the 
green-yellow  spotted-red  blotched  pattern.  There  were,  also,  9  cases  of 
spontaneous  development  of  j^ellow  that  were  not  carried  on  in  suc- 
cessive leaves,  and  with  the  exception  of  1  case  were  confined  to  but 
one  or  two  leaves.  These  yellow  blotches  were  large,  irregular-shaped 
pure-yellow  areas  covering  from  one-eighth  to  one-fourth  the  entire 
area  of  a  leaf.  The  locations  and  relative  sizes  of  these  yellow  blotches 
in  the  leaves  of  one  plant  are  shown  in  text-figure  1,  the  shaded  protions 
of  which  indicate  yellow  areas.  All  other  leaves  were  pure  green  and 
the  branches  produced  in  the  axils  of  the  yellow-blotched  leaves  were 
pure  green. 


Text-figure  1. — Position  of  the  yellow  areas  that  developed  spontaneously  in  these 

leaves  of  plant  11714221. 

A  study  of  pedigrees  reveals  the  interesting  fact  that  6  of  the  plants 
with  spontaneous  yellow  all  descended  from  a  branch  on  plant  11714. 
The  full  record  of  this  clone  is  given  in  table  13,  but  the  summary  of 
the  pedigrees  of  the  particular  line  of  descent  is  here  given  in  table  6. 

The  pedigree  numbers  enable  one  to  trace  relationship  quite  readil3^ 
In  this  case  we  note  that  plant  1171  gave  a  bud  variation  with  loss  of 
yellow.  Plant  11714  was  grown  as  a  chimera  with  about  one-half  pure 
green.  From  the  green  part  two  cuttings  were  made  for  plants  117142 
and  117144,  both  of  which  remained  constant  for  loss  of  yellow.  In 
September  1913,  three  cuttings  were  made  and  from  two  of  the  plants 
grown  three  more  cuttings  were  taken  in  the  following  spring.  From 
September  1912  until  August  1914  all  plants  grown  in  these  lines  of 
descent  were  constant  for  loss  of  yellow,  then  in  one  season  the  spon- 
taneous development  of  yellow  occured  in  6  closely  related  plants. 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS. 


31 


Table  6. — Pedigree  of  6  plants  giving  spontaneous  yellow. 


Time. 


Plant. 


Pattern. 


Record. 


Apr.    1912  to  Sept.  1914 
Sept.  1912  to  Sept.  1913 

Summer  1913 


Sept.  1913  to  Sept.  1914 


Summer  1914 . 


117 Green  -  yellow  -  red 

blotched 

117,  1 do 


117  14»  /••••do 

\Green-red  blotched . 

117,  142 do 

117,  144 do 

117,142,1.. do 


117,142,2 do. 

117,144,1 do. 

117.142.11. do. 

117.142.12. do. 

[117,142,21..] do. 


Constant. 

Bud   variation  to  green- 
red  blotched. 
Constant. 
Constant  for  loss  of  yellow. 

Do. 

Do. 
Spontaneous    yellow     in 
August. 

Do. 

Do. 

Do. 

Do. 

Do. 


'Chimera. 

Including  the  cases  of  spontaneous  appearance  of  yellow  with  sec- 
torial bud  variations,  the  ratio  of  frequency  was  1  to  1,120.  The  ratio 
for  the  sectorial  bud  variations  alone  was  about  1  to  2,570. 

Plants  with  pattern  yellow-green-red  blotched. — All  of  the  plants  grown 
with  this  pattern  (fig.  6)  belonged  to  clone  11,  and  all,  excepting  3,  de- 
scended from  the  plant  1171,  which  produced  a  series  of  branches  with 
this  pattern,  as  already  shown  in  diagram  1.  The  record  of  these 
plants  is  given  later  in  table  13,  but  may  be  summarized  here  as  given 
in  table  7. 

Table  7. — Summary  of  plants  loith  yellow-green-red  blotched  pattern  {fig.  6),  all 

of  clone  11. 


Yellow-green-red  blotched 

p.  .  fyellow-green-red  blotched. . 

^yellow-red  blotched 

/^f  Jyellow-green-red  blotched. . 

Chimera  <  j  ui  *  u    i 

1  green-red  blotched 

CK  fyellow-green-red  blotched. . 

\yellow-green 

Total 


o 

a 


o 


29 

8 


a 


O 


17 
5 


41     24 


a 

"■+3 

3 
o 

3 


M  a 
a  o 


Bud  variations. 


•n 

(D 

O  -C 
-I 


9       3 


13 


1^ 

cl  aj  o 
g  +^    O 

Op 


a;  ^ 
(U    o 


10 


a 

i-i 
60 
I 

"53 


32  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

In  several  instances  cuttings  were  so  made  that  the  resulting  plants 
possessed  two  patterns.  These  plants  are  included  here  in  respect  to 
the  behavior  of  the  part  with  pattern  yellow-green-red  blotched. 

All  of  the  plants  grown  during  the  winter  showed  more  or  less 
increase  of  green,  but  as  long  as  the  yellow  was  present  as  a  definite 
central  area  they  were  classed  as  constant.  There  were,  for  example, 
just  such  differences  in  development  of  yellow  as  is  seen  in  figures 
14  and  14a,  the  former  representing  the  usual  condition  during  winter 
and  the  latter  the  development  of  yellow  during  the  summer.  Four 
plants  fluctuated  in  marked  degree,  giving  mixed  and  irregular  patterns 
with  many  leaves  in  which  there  was  much  green. 

One  case  of  bud  variation  was  concerned  with  loss  of  epidermal 
red,  giving  the  type  yellow-green  (fig.  14).  The  other  instances  gave 
3  cases  of  extreme  development  of  yellow,  10  cases  of  entire  loss  of 
yellow,  and  1  case  of  change  to  the  yellow  spotted  condition.  The 
latter,  however,  occurred  on  a  plant  with  also  a  bud  variation  to  pure 
green-red  blotched.  The  ratio  of  frequency  for  bud  variation  in  this 
group  was  about  1  to  540. 

Plants  with  pattern  green-yelloiv-solid  red. — In  respect  to  the  green 
and  yellow  this  pattern  (fig.  8)  is  identical  with  that  of  green-yellow- 
red  blotched.  It  differs  in  having  a  solid-red  instead  of  a  red-blotched 
epidermis.  The  54  plants  grown  with  this  pattern  remained  free  from 
any  noticeable  variations  in  respect  to  the  solid  red  epidermis,  except 
those  that  were  bud  variations.  Frequently  a  leaf  appeared  with  a 
few  small  areas  in  which  the  red  of  the  epidermis  was  absent,  but  these 
were  rather  isolated.  There  was  some  degree  of  fluctuation  in  the 
relative  amounts  of  green  and  yellow,  with  a  tendency  for  plants  to  be 
greener  in  winter  and  yellower  in  summer.  On  account  of  the  soUd  red, 
it  was  more  difficult  to  judge  these  fluctuations  than  in  plants  with 
red  blotched  or  with  non-red  epidermis,  hence  attention  was  chiefly 
directed  to  the  condition  of  the  epidermis.  In  making  cuttings,  plants 
most  constant  and  typical  for  the  green-yellow  condition  were,  however, 
selected. 

Of  the  10  cases  of  bud  variation,  4  gave  complete  loss  of  yellow, 
2  gave  a  reversal  of  the  relative  position  of  green  and  yellow,  and  1 
gave  extreme  development  of  yellow.  Only  3  cases  involved  variation 
in  the  amount  and  distribution  of  red;  1  was  a  complete  loss  and  2  gave 
the  pattern  described  above  as  green-yellow-solid  red  upper  center, 
a  type  which  is  an  interesting  intermediate  between  no  red  and  solid  red. 
The  ratio  of  frequency  for  all  bud  variations  for  the  group  is  1  to  1,080. 

Plants  with  green-solid  red  pattern. — This  pattern  (fig.  9)  first  ap- 
peared during  the  summer  of  1913  as  a  bud  variation  on  a  plant  of 
pattern  green-red  blotched.  From  this  branch  cuttings  were  taken  for 
6  plants  grown  during  the  summer  of  1914.  All  of  these  remained  con- 
stant for  loss  of  yellow  and  for  a  solid-red  epidermis,  except  1  plant. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS. 


33 


upon  which  a  sectorial  variation  in  one  small  branch  gave  the  pattern 
green. 

With  respect  to  the  red  epidermis,  patterns  green-yellow-solid  red 
(fig.  8)  and  green-solid  red  are  identical,  and  it  is  noteworthy  that  in  the 
62  plants  of  these  two  patterns  which  were  grown  to  maturity  there 
were  no  noticeable  fluctuations  and  but  4  cases  of  bud  variations  involv- 
ing the  red.  For  the  red  epidermis  the  ratio  of  frequency  of  bud  varia- 
tions was  1  to  3,100. 

Table  8. — Summary  of  plcmls  ivilh  solid  red  -patterns,  clone  S. 


Green-yellow- 
solid  red. 

Green-solid 
red. 

Total  number  of  plants 

54 

51 

9 

1 
2 

4 
1 

8 
7 
1 

1 

Constant  for  solid  red 

Number  of  plants  with  bud  variations. .  . 
Bud  variations: 

To  vellow-solid  red 

To  vellow-green-solid  red 

To  green-yellow-solid  red  upper  center . 
To  green-solid  red  .  . 

To  green-yellow 

To  green 

Plants  with  pattern  green-yellow-solid  red  upper  center. — This  bright 
and  attractive  pattern  (fig.  10)  first  developed  as  a  sectorial  bud  vari- 
ation in  the  winter  of  1912  on  a  plant  otherwise  uniform  and  constant 
to  type  green-yellow-solid  red.  During  the  summer  of  1913,  the  plant 
grew  vigorously  and  numerous  branches  developed  from  the  part  hav- 
ing this  new  pattern,  all  of  which  were  constant  and  uniform  for  the 
new  type  and  which  were  in  conspicuous  contrast  to  the  rest  of  the  plant. 

In  the  autumn  of  1913  cuttings  were  made  from  these  branches  for 
5  plants,  which  were  grown  until  the  autumn  of  1914.  Four  of  these 
remained  quite  constant,  although  they  were  much  greener  in  winter. 
One  plant  became  gradually  greener  during  the  early  part  of  winter 
until  it  was  apparently  pure  green,  giving  type  green-solid  red  upper 
center.  The  upper  surface  of  a  leaf  of  this  plant  painted  in  January 
is  shown  in  figure  15.  Toward  spring  the  new  leaves  produced  by  this 
plant  became  quite  laciniate,  but  during  the  following  summer  the 
leaves  produced  were  entire  and  strongly  tinged  with  yellow.  During 
the  summer  of  1914,  two  plants  from  cuttings  of  one  of  the  plants 
constant  for  the  type  remained  true  to  that  pattern. 

No  bud  variations  appeared  in  any  of  the  7  plants  and  no  noticeable 
fluctuations  in  the  amount  and  distribution  of  the  red ;  yet  there  was 
no  plant  that  did  not  show  at  some  time  a  few  leaves  with  tiny  red  spots 
scattered  on  the  lower  surface,  much  as  is  shown  in  figure  10a. 

This  pattern  also  appeared  late  in  the  summer  of  1914  as  a  sectorial 
variation  on  a  plant  which  during  the  summer  had  been  constant  and 
uniform  for  type  green-yellow-solid  red.     A  cutting  was  made  from  this 


34  ESTABLISHMENT    OF   VARIETIES    IX    COLEUS 

branch,  and  the  young  plants  grown  from  it  are  at  the  present  writing 
(December  10,  1914)  nearly  devoid  of  yellow,  but  have  the  change  in 
pattern  for  red  as  a  clear-cut  sectorial  variation.  Both  surfaces  of  a 
single  leaf  are  shown  in  figures  24  and  24a.  In  figure  24  the  loss  of 
epidermal  red  on  the  lower  surface  of  half  of  the  leaf  illustrates  very  well 
the  definiteness  with  which  color  variations  in  Coleus  appear.  The 
upper  surface  of  the  corresponding  half  of  this  leaf  is  shown  in  figure 
24a,  with  the  decrease  of  red  about  the  margin.  Such  differences  are 
usually  seen  in  a  series  of  leaves  in  the  same  row  and  in  the  branches 
that  develop  in  the  axils  of  such  leaves,  gi\dng  a  marked  degree  of 
sectorial  symmetry  to  the  distribution  of  pigmentation,  a  condition  also 
well  illustrated  with  reference  to  green  and  yellow  in  figure  21. 

Plants  with  pattern  yellow-green-solid  red. — This  pattern  (fig.  11)  first 
appeared  during  the  summer  of  1913  as  a  variation  complete  for  a  single 
lateral  branch  of  a  plant  with  green-yellow-solid  red.  This  branch  was 
removed  for  a  cutting,  but  died  soon  after  it  was  rooted  and  placed  in 
a  pot.  Early  in  the  spring  of  1914  this  pattern  appeared  as  a  sectorial 
variation  in  the  main  axis  of  a  plant  having  green-yellow-solid  red.  This 
plant  grew  vigorously,  giving  a  large,  bushj^  plant  with  the  two  types 
of  foliage  distinct  and  constant  on  the  different  branches.  Numerous 
cuttings  have  been  made  to  test  the  vegetative  constancy  of  this  type. 

Plants  with  -pattern  green-yelloiv. — Three  plants  of  this  type  (fig.  12) 
were  grown  from  September  1913  until  October  1914  and  4  were  grown 
during  the  summer  of  1914.  All  of  these  remained  quite  constant  for 
the  loss  of  epidermal  red.  They  were  much  less  uniform  in  regard  to 
the  relative  amounts  of  green  and  yellow,  one  plant  possessing  a  branch 
that  was  quite  green.  There  was  also  a  strong  tendency  among  the 
leaves  on  one  plant  to  show  somewhat  irregular  distribution  by  green 
and  yellow,  as  is  shown  in  figure  12. 

Plants  with  pattern  green  and  pattern  green-yellow  spotted. — The  loss 
of  epidermal  red  occurred  as  a  sectorial  bud  variation  during  the  late 
summer  of  1913  on  a  plant  that  had  fluctuated  from  type  green-yellow- 
red  blotched  to  type  green-yellow  spotted-red  blotched.  The  develop- 
ment of  yellow  was,  however,  very  faint,  so  that  the  bud  variation 
gave  a  leaf  pattern  that  was  almost  pure  green.  This  plant  was 
taken  up  and  grown  in  a  pot  during  the  winter  of  1913-14  and  during 
the  following  summer  again  grown  out  of  doors.  All  branches  on  the 
two  parts  were  quite  constant  in  respect  to  presence  and  absence  of  the 
epidermal  red,  but  there  were  traces  of  red  coloration  in  the  sub-epider- 
mal tissues  which,  as  shown  in  figure  13,  were  almost  entirel}^  confined 
to  the  vascular  tissues.  There  was  more  or  less  fluctuation  on  the  entire 
plant  in  the  appearance  of  yellow,  but  no  decided  development  of  it. 

In  January  1913  a  cutting  was  made  from  the  part  of  this  plant 
which  showed  the  bud  variation.  The  plant  grew  vigorously  and  was 
during  the  winter  mostly  free  from  yellow.     In  April,  5  cuttings  were 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS. 


35 


made  for  plants.  During  the  summer  of  1914  the  G  plants  remained 
constant  for  loss  of  epidermal  red,  but  bore  some  leaves  with  yellow 
spots.  There  was,  however,  a  rather  weak  development  of  yellow  and 
from  a  short  distance  the  plants  appeared  to  be  pure  green.  The 
fluctuations  in  the  development  of  green  and  yellow  are  quite  like 
those  in  other  patterns.  The  pattern  is  of  special  interest  in  regard 
to  the  development  of  red  pigmentation  in  the  subepidermal  tissues, 
especially  in  the  vascular  elements  giving  a  reticulated  effect  well 
shown  in  figure  13,  and  also  seen  in  figure  14a.  This  condition 
also  prevails  in  the  red  blotched  types,  but  is  more  or  less  obscured 
by  the  more  conspicuous  epidermal  coloration. 

During  the  summer  of  1914  there  was  no  noticeable  variation  in 
the  amount  of  red  in  subepidermal  tissues.  Cuttings  made  in  the 
autumn  of  that  year  for  a  new  generation  exhibited  during  the  winter 
marked  variations  in  this  respect.  The  summaries  given  in  this  paper 
do  not  include  the  generation  to  which  these  plants  belong,  but  the 
behavior  of  this  particular  set  of  plants  can  be  included  here.     Figure 

Table  9. — Summary  of  plants  with  non  red  epidermis. 


Pattern. 


Green-yellow 

Green  and  green-yellow  spotted . 
Yellow-green 


Total 
plants. 


7 
7 
4 


Constant 
for  non  red 
epidermis. 


7 
7 
4 


Plants 
giving  bud 
variations. 


0 
0 
0 


13d  gives  a  leaf  painted  on  February  2,  1915,  showing  the  development 
of  red  in  the  internal  tissues.  Free-hand  sections  of  such  leaves  indi- 
cated that  the  epidermal  cells  are  non-red.  The  coloration  appears 
dull,  as  if  glazed  over  rather  than  velvety  as  in  the  epidermal  colora- 
tion, a  contrast  due  largely  to  the  coloration  of  the  trichomes  of 
the  epidermis  and  which  the  reproductions  do  not  adequately  show. 

Plants  with  'pattern  yellow-green. — This  pattern  (fig.  14)  identical  with 
that  of  type  yellow-green-red  blotched  except  for  the  loss  of  epidermal 
red,  appeared  as  a  sectorial  bud  variation  late  in  the  summer  of  1913. 
The  sporting  branch  was  used  as  a  cutting,  from  which  a  large  plant 
grew  during  the  winter  of  1913  and  the  following  summer.  In  the 
winter  there  was  an  increase  of  green,  but  throughout  its  growth  the 
part  with  pattern  yellow-green  remained  constant  in  respect  to  the  loss 
of  epidermal  red. 

Three  plants  of  a  new  generation  were  grown  during  the  summer  of 
1914,  and  these  remained  constant  and  uniform  for  the  loss  of  epidermal 
red.  In  regard  to  the  relative  positions  of  green  and  yellow,  the  plants 
were  quite  constant,  but  there  was  a  strong  tendency  for  green  to 
increase  in  winter  and  decrease  in  summer,  giving  such  differences  as 
are  shown  in  figures  14  and  14a. 


36  ESTABLISHMENT    OF   VARIETIES   IN    COLEUS 

The  figures  13,  a,  b,  and  c,  show  the  three  leaves  growing  in  the 
rank  above  the  leaf  shown  in  figure  13d  and  ilkistrate  the  increase  of 
red  which  very  plainly  occurs  as  the  leaves  mature.  In  the  red- 
blotched  and  solid  red  patterns  an  increase  in  the  total  amount  of  red 
pigmentation  must,  it  would  seem,  also  occur  as  the  leaves  enlarge. 

As  the  last  four  patterns,  green-yellow,  green,  green-yellow  spotted,  and 
yellow-green  are  alike  in  respect  to  loss  of  epidermal  red,  they  may  be 
grouped  in  this  respect.  It  is  noteworthy  that  there  was  no  case  of  a 
development  of  epidermal  red.  No  plant  was  free  of  some  red  colora- 
tion in  stems  and  in  vascular  strands  of  the  leaves,  as  especially  well 
shown  in  figures  13  and  14,  There  were  cases  where  the  coloration 
seemed  to  spread  out  near  the  ends  of  the  vascular  strands,  but  the 
appearance  was  not  the  same  as  that  of  the  blotches.  The  number  of 
plants  of  these  patterns  grown  thus  far  is  small  and  their  behavior  is 
not  taken  as  fully  indicative  of  the  possible  variations  that  may  appear 
in  future  cultures. 


From  the  summaries  of  patterns  given  above  it  is  quite  clear 
that  the  various  types  noted  (with  the  exception  of  pattern  green-solid 
red  upper  center)  have  been  kept  quite  constant  by  a  selection  of  the 
parent  plants  to  be  used  in  vegetative  propagation,  and  that  every  new 
type  of  pattern  (excepting  the  one)  that  arose  either  by  fluctuating 
variation  or  by  bud  variation  can  be  propagated  as  a  vegetative  type. 
It  is  highly  possible  that  finer  distinctions  could  be  made  in  regard  to 
pattern  types,  especially  inside  of  the  rather  comprehensive  groups 
classed  as  red  blotched,  as  yellow  bordered,  and  as  yellow  spotted,  among 
which  there  were  many  variations  that  gave  all  degrees  of  gradation  to 
or  even  into  a  different  pattern.  The  writer  wishes  to  state  that  the 
keeping  of  records  satisfactory  to  him  was  no  simple  matter,  even  for 
the  pattern  classes  as  determined. 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS. 


37 


RANGE  OF  THE  VARIATIONS. 

At  this  point  the  data  already  presented  may  be  summarized  and  anal- 
yzed in  reference  to  the  range  or  extent  of  the  variations  as  a  whole. 

There  have  already  been  described  15  different  color  patterns  that 
arose  by  bud  variation,  1  very  decided  color  pattern  that  arose  solely 
as  a  fluctuating  variation,  and  the  laciniate  type  of  leaf.  Several  of 
the  principal  types  arose  also  by  fluctuating  variations.  In  respect  to 
the  relative  amounts  of  green  and  yellow  there  are  the  two  extremes: 
(a)  almost  pure  development  of  yellow  (fig.  1),  and  (b)  pure  green 
(figs.  5  and  9),  with  almost  every  possible  gradation  between.  Of  the 
epidermal  red  there  are  the  extremes  solid  red  and  no  red  with  the 
intermediate  red-blotched  type  (including  wide  variations),  and  the  type 
solid  red  upper  center  as  another  intermediate.  In  respect  to  the  relative 
positions  of  the  green  and  yellow  there  are  the  extremes:  (a)  green  center 
w^ith  yellow  border  (fig.  2)  and  (6)  yellow  center  with  green  border  (fig.  6) . 
Between  these  the  irregular  patterns  present  numerous  intermediates. 


,1.  Yellow-red 
blotched 


/A-    Green- 
/yellowspotled 
red  blotched 


Spontaneous  yellow 

7 2or4andE-----5and7 


8a.Greer>-yellowspcrtted-sclid  ned 
13a.  Greeri -yellow  spotted 


Z.Grccn-yCllow-  . 
red   blolcheci 


.  5.  Green-red 
blotched 


6.  Yellow- 
green-  red 
blotched 


8.  Green- 


yellow- 
Solid  red 


Spontaneous  yellow 

*•  ^5  and  t 

2— -4  and    E Send? 

9.  Green- sol  id  red         '^® 

13.  Green I3e 

I. 

4 
5. 

14.  Yellow-green 

9.  Green -sol  id  red 

10.  Green-yellow-solid  red  upper  center' 

11.  Yellovy-green-solid  red 
12. 

16.  Yellow-solid  red 


.i5eryJ7 


12.  Green-yellow 
Diagram  3. — Derivations  of  color  patterns. 


Diagram  3  gives  a  graphic  representation  of  the  extent  of  variations 
in  each  pattern  and  shows  the  derivation  of  types  and  the  appearance 
of  the  same  types  of  pattern  as  variations  from  quite  different  patterns. 


38  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

In  diagram  3  the  numbers  (except  13  a)  refer  to  figures  illustrating 
the  types  of  color  pattern  or  leaf-shape.  The  name  of  the  pattern  is 
written  in  full  only  in  the  line  of  descent  when  it  first  appeared.  A 
continuous  Une  indicates  origin  by  bud  variation,  while  a  broken  line 
indicates  fluctuation. 

From  the  original  pattern  of  green-yellow-red  blotched  there  arose 
directlj'  6  different  patterns  (see  tables  1  and  2  and  diagram  3)  each 
involving  a  single  marked  variation.  For  the  development  of  any 
other  pattern  thus  far  produced,  excepting  the  green-yellow-solid  red 
upper  center  pattern  a  second  change  is  necessary.  On  any  of  these  6 
derived  patterns  a  further  single  change  may  give  a  new  pattern  or 
produce  a  pattern  already  realized.  The  bud  variations  in  the  derived 
pattern  green-yellow  spotted-red  blotched  (fig.  4)  illustrate  this  point. 
Increase  of  yellow  gave  yellow-red  blotched  (fig.  1)  and  loss  of  yellow 
gave  green-red  blotched  (fig.  5),  both  patterns  previously  derived,  and 
also  a  return  to  the  original  type  of  green-yellow-red  blotched  (fig.  2). 
In  all  these  the  red-blotched  condition  of  the  epidermis  is  much  the 
same.  Changes  in  the  epidermal  coloration,  however,  give  new  pat- 
terns. The  appearance  of  solid  red  gives  a  slightly  different  pattern 
than  that  of  green-yellow-solid  red.  The  pattern  green-yellow  spotted 
appeared  as  a  bud  variation  by  a  loss  of  the  epidermal  red. 

The  green-red  blotched  (fig.  5)  pattern  gave  opportunity  for  new 
patterns  by  the  same  changes  in  epidermal  red  which  have  previously 
appeared.  These  are  realized  in  green-solid  red  (fig.  9)  and  green-no  red 
(fig.  13).  There  is  also  chance  for  reappearance  of  yellow  to  give  return 
to  old  types  or  possibly  to  new  types.  Of  these  only  one  appeared,  and 
this  was  the  pattern  green-yellow  spotted-red  blotched  (fig.  4).  The  cases 
of  spontaneous  appearance  of  yellow  did  not  result  in  a  definite  pattern. 
In  this  line  of  descent  there  also  developed  the  character  of  laciniate 
shape  of  the  leaf,  with  its  marked  periodicity  of  expression. 

The  changes  in  the  pattern  yellow-green-red  blotched  (fig.  6)  which 
involved  amounts  of  green  and  yellow  gave  no  new  patterns.  The 
loss  of  red,  however,  gave  a  new  pattern  yellow-green  (fig.  14).  At 
least  3  possible  bud  sports  did  not  appear  in  this  pattern:  (1)  changes 
producing  a  reversal  of  green  and  yellow  giving  return  to  the  parent 
type;  (2)  a  solid-red  (fig.  11);  or  (3)  solid  red  upper  center  (fig.  10). 

In  the  plants  with  green-yellow-solid  red  (fig.  8),  a  loss  of  yellow  gave 
the  same  pattern  that  was  produced  by  gain  of  soUd  red  from  pattern 
green-red  blotched.  The  two  changes  involved  are  identical,  but  occurred 
in  reversed  order.  Reversal  of  the  positions  of  green  and  yellow,  a 
change  identical  with  that  giving  the  yellow-green-red  blotched  pattern, 
gave  a  different  pattern  because  the  tissues  were  overlaid  by  solid  red. 
The  same  is  true  of  the  loss  of  green.  Loss  of  the  epidermal  red  on  the 
lower  surface  and  about  the  margin  of  the  upper  surface  gave  a  pattern 
green-yellow-solid  red  upper  center  (fig.  10)  that  has  not  appeared  else- 
where and  is  really  the  only  new  variation  that  appeared  in  this  pattern. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  39 

Plants  grown  from  pattern  green-yellow-red  blotched  (fig.  2)  derived 
from  green-yelloiv  spotted-red  blotched  (fig.  4)  gave  again  the  same  pat- 
terns that  were  directly  derived  from  that  pattern. 

A  general  review  of  the  entire  series  of  variations  shows  that  in 
respect  to  the  relative  positions  and  total  amounts  of  green  and  yellow, 
and  the  total  amount  of  red  in  the  epidermis,  the  extremes  of  develop- 
ment possible  are  reahzed,  with,  also,  the  appearance  of  a  series  of  inter- 
mediate types.  Judging  the  variations  in  any  one  pattern  by  the 
range  of  bud  variations  that  have  thus  far  developed,  it  appears  that 
any  pattern  tested  in  considerable  numbers  gives  by  bud  variation  the 
entire  range  of  changes  possible. 

FREQUENCY  OF  THE  BUD  VARIATIONS. 

With  the  list  of  the  tj^pes  of  variations  given  in  table  1  there  is  also 
given  the  data  as  to  the  number  of  times  each  occurred  and  the  total 
number  of  plants  involved  (not  including  plants  of  patterns  which  did 
not  produce  the  particular  variation) .  The  various  tables  present  the 
details  of  their  data,  which  may  be  now  summarized  under  the  main 
types  of  changes  outlined  in  table  1. 

A.    CHANGES    INVOLVING    YELLOW    AND    GREEN. 
I.  Increase  of  Yellow  and  Decrease  of  Green. 

1.  The  almost  complete  loss  of  green  with  increase  of  yellow  occurred 
as  a  bud  variation  12  times  in  a  total  of  630  plants  (not  including  plants 
of  patterns  which  did  not  give  this  variation).  It  was  derived  most 
frequently  from  patterns  green-yellow-red  blotched  (fig.  2),  yellow-green- 
red  blotched  (fig.  6),  and  green-yellow-solid  red  (fig.  8).  It  developed  once 
from  pattern  green-yellow  spotted-red  blotched  (fig.  4).  The  pattern 
was  not  realized  uniformly  on  any  plant  as  a  fluctuating  variation.  On 
a  few  plants  of  the  pattern  green-yellow-red  blotched  which  developed 
mixed  patterns  with  a  marked  increase  of  yellow  during  the  summer, 
some  leaves  approached  this  pattern.  One  of  the  most  marked  of 
these  is  shown  in  figure  20. 

2.  It  will  be  remembered  that  the  green-yellow-red  blotched  pattern 
was  borne  by  the  parent  plants  and  that  all  other  patterns  were  derived 
directly  or  indirectly  from  this.  The  return  to  this  pattern  occurred 
as  a  bud  variation  from  type  green-yellow  spotted-red  blotched  in  4 
instances  on  a  total  of  266  plants  and  also  as  fluctuating  variations, 
especially  in  clone  14,  as  shown  in  table  3. 

3.  The  sudden  appearance  of  scattered  areas  of  yellow  in  single 
branches  of  plants  otherwise  having  no  yellow  occurred  but  twice. 
The  same  type  green-yellow  spotted-red  blotched,  however,  appeared 
quite  gradually  for  entire  plants  in  38  cases  out  of  the  90  plants  which 
were  grown  for  the  pattern  green-red  blotched. 


40  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

4.  The  development  of  rather  large  conspicuous  areas  of  yellow  in 
one  or  more  leaves  of  a  plant  otherwise  pure  green  occurred  9  times. 
In  one  case  three  leaves  of  the  same  branch  showed  yellow  areas  that 
arose  in  this  apparently  spontaneous  manner  (see  text-fig.  1). 

Summary. — There  were  27  cases  of  bud  variation  giving  increase  of 
yellow  and  involving  directlj^  788  plants.  There  was  opportunity 
for  increase  of  yellow  to  occur  in  all  plants  grown,  except  those  of  the 
pattern  yellow-red  blotched,  and  even  in  these,  10  of  the  11  were  grown 
as  chimeras  with  at  least  half  of  the  plant  green.  In  computing  a 
final  ratio  for  the  frequency  of  bud  variations  giving  increase  of  yellow, 
we  may  use  all  but  6  of  the  plants  grown.  The  ratio  of  frequency  on 
this  basis  is  1  to  6,130. 

II.  Decrease  of  Yellow  and  Increase  of  Green. 

1 .  Pattern  green-yellow  spotted-red  blotched  was  produced  from  green- 
yellow-red  blotched  and  yelloiv-green-red  blotched  in  9  instances  on  a  total 
of  378  plants.  In  4  other  cases  the  change  to  yellow  spotted  affected 
single  branches,  and  although  marked  for  a  time  after  the  first  appear- 
ance, later  fluctuated  to  the  parent  type  green-yellow-red  spotted,  for 
which  the  plants  became  quite  uniform. 

2.  Pattern  green-red  blotched  with  complete  loss  of  yellow  occurred  on 
plants  with  green-yellow-red  blotched,  with  green-yellow  spotted-red  blotched 
(entire  and  laciniate),  and  with  yellow-green-red  blotched  patterns  in 
37  instances  on  a  total  of  644  plants.  The  same  change  gave  pattern 
green-solid  red  4  times  on  54  plants  of  pattern  green-yellow-solid  red. 

Summary. — Bud  variations  producing  increase  of  green  occurred  50 
times.  The  total  plants  grown  with  more  or  less  j^ellow  were  740. 
The  ratio  of  frequency  for  loss  of  green  by  bud  variation  was  1  to  2,960. 

III.  Reversal  of  the  Relative  Positions  of  Green  and  Yellow. 

1 .  This  reversal  has  only  occurred  in  patterns  with  the  yellow  at  the 
border  of  the  leaf,  giving  yellow-green-red  blotched  (fig.  6)  from  green- 
yellow-red  blotched  (fig.  2)  and  yellow-green-solid  red  (fig.  11)  from  green- 
yellow-solid  red  (fig.  8)  in  a  total  of  8  instances  on  391  plants.  It  is  also 
possible  for  a  reversal  to  occur  in  any  other  patterns  having  a  distinct 
border  of  green  or  yellow.  The  total  of  such  plants  is  450,  which  gives 
1  to  11,250  as  the  ratio  of  frequency  for  this  change. 

B.    CHANGES  INVOLVING  THE  EPIDERMAL  RED. 
I.  Increase  of  Epidermal  Red. 

Eight  instances  of  bud  variations  giving  solid  red  occurred  in  red 
blotched  patterns  involving  directly  a  total  of  625  plants.  None  of 
the  41  plants  of  the  pattern  yellow-green-red  blotched  gave  this  variation. 
Increase  of  red  was  possible  in  all  except  the  solid-red  patterns  (62 
plants  in  all).     The  ratio  for  this  change  was  1  to  19,250. 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS. 


41 


II.  Decrease  of  Epidekmal  Red. 

Bud  variations  of  this  sort  can  be  graded  as  follows:  an  almost  com- 
plete loss  of  red  on  the  under  surface  and  about  the  upper  margin, 
which  occurred  2  times,  and  an  apparently  complete  loss  of  epidermal 
red  in  patterns  with  spotted  or  with  solid  red  epidermis,  which  appeared 
in  19  instances.  A  total  of  815  plants  were  grown  of  patterns  having 
some  degree  of  red  in  the  epidermis.  The  ratio  for  complete  loss  of  red 
was  1  to  8,580  and  for  all  cases  of  decrease  of  red  it  was  1  to  7,760. 

C.    CHANGES  INVOLVING  LEAF-SHAPE. 

The  appearance  of  the  laciniate  leaf-shape  as  a  fluctuating  variation 
which  marked  periodicity  of  development  occurred  13  times.  (The 
bud  variation  giving  this  type  late  in  1914  is  not  included.)  The 
total  number  of  plants  grown  with  entire  leaves  was  765,  hence  the  ratio 
on  the  basis  used  hitherto  was  1  to  11,770.  It  seems,  however,  that 
this  basis  hardly  affords  the  same  degree  of  accuracy  for  comparison  as 
it  does  between  the  different  bud  variations  in  color.  Here  the  change 
appeared  in  an  entire  plant  (except  one  plant  grown  during  winter 
of  1914-15  and  not  included  in  these  computations),  but  as  several  of 
these  were  from  a  same  immediate  parent,  it  may  be  that  the  change 
really  arose  as  a  bud  variation,  with,  however,  a  delayed  effect. 

Table  10. — Frequency  of  changes  giving  the  different  types. 


Type  of  change. 

Plants. 

Frequency. 

Ratio. 

Increase  of  yellow  and  decrease  of  green  .... 

Decrease  of  yellow  and  increase  of  green 

Reversal  of  positions  of  green  and  yellow .... 

Increase  of  epidermal  red  to  solid  red 

Decrease  of  epidermal  red,  complete  loss .... 

Decrease  of  epidermal  red,  all  cases 

Appearance  of  the  laciniate  character 

Entire  leaf  from  laciniate  leaf 

827 
740 
450 
770 
815 
815 
765 
68 

27 

50 

8 

8 

19 

21 

13 

1 

6,1.30 

2,960 

11,250 

19,250 

8,580 

7,760 

11,770 

13,600 

Of  the  68  plants  grown  with  the  habit  of  producing  laciniate  leaves, 
a  single  case  of  persistent  change  to  the  entire  leaf-shape  appeared. 
There  were  also  3  cases  of  clear-cut  bud  variations  involving  color 
changes  in  these  plants. 

SUMMARY  AND  COMPARISONS. 

For  the  purpose  of  comparison,  the  ratios  showing  the  frequency 
with  which  these  different  types  of  changes  appear  are  brought  together 
in  table  10.  In  deriving  these  ratios  the  total  number  of  plants  in  which 
there  is  possibiHty  for  the  change  to  occur  has  been  considered. 

These  data  indicate  the  tendencies  of  the  bud  variations  and  give  a 
clew  to  the  behavior  of  the  characters  in  question.  In  the  bud  vari- 
ations, decrease  of  yellow  occurred  twice  as  often  as  the  increase  of 
yellow.     Likewise,  the  loss  of  red  occurred  2.2  times  as  often  as  the 


42 


ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 


increase  of  red.  If  we  consider  that  the  abihty  to  produce  yellow  and 
red  are  the  more  recently  acquired  characters  of  the  cells,  these  data 
would  indicate  a  tendency  toward  loss  rather  than  gain  of  these 
characters. 

A  summary  of  the  data  regarding  the  degree  of  constancy  of  the 
various  patterns  and  the  nature  of  the  variations  which  they  exhibit 
is  of  further  interest  in  a  consideration  of  the  tendencies  of  the  vari- 
ations. In  comparing  bud  variations  which  originate  in  a  bud  the 
comparison  on  the  basis  of  the  total  buds  produced  seems  quite  ade- 
quate. The  comparison  of  fluctuating  variations  requires  a  different 
treatment.  On  plants  with  irregular  and  mixed  patterns  it  is  not 
practicable,  if  possible,  to  attempt  a  statistical  determination  of  the 
fluctuating  branches.     Only  in  few  cases  when  such   changes  were 

Table  1 1 . — Summary  of  changes  occurring  in  the  principal  patterns. 


Green-  Yellow- 
yellow-  green- 
red  red 
blotched,  blotched. 


Green- 
yellow 

spotted- 
red 

blotched. 


Plants 

with 

laciniate 

leaves. 


Green- 
red 
blotched. 


Total  number  plants '  337 

Plants  constant  for  green  and  yellow.  .  .  218 

Percentage  of  constant  plants 65 

Changes  in  yellow: 
Increase: 

Frequency 7 

Ratio  of  frequency 9,630 

Decrease : 

Frequency 32 

Ratio  of  frequency 2,100 

Reversal :  j 

Frequency 6 

Ratio  of  frequency ■  11,230 

Total  frequency 45 

Ratio  of  frequency  for  all  bud  variations .  1 ,490 


41 
24 
59 


3 

2,730 

11 

740 


14 
590 


198 

126 

63 


4 
9,900 

2 
19,800 


6 
0,600 


68 
0 
0 


1 

13,600 

1 
13,600 


90 
31 
34 


11 
1,6.30 


2 
6,800 


11 
1,630 


Umited  to  a  branch  could  there  by  any  degree  of  accuracy.  Further- 
more, fluctuations  in  number  and  size  of  the  blotches  of  epidermal  red, 
although  frequent  and  somewhat  persistent,  were  not  recorded.  As 
long  as  the  pattern  was  blotched  the  plants  were  grouped  together  and 
changes  to  solid  red  or  to  no-red  for  considerable  areas  of  a  leaf  were 
not  considered  as  a  bud  variation  unless  a  series  of  leaves  showed  that 
the  change  was  sectorial  for  a  stem.  For  this  reason  the  data  given  in 
table  11  are  summarized  for  fluctuations  and  bud  variations  involving 
yellow  and  green  in  patterns  with  red  blotched  epidermis. 

The  percentage  of  constant  plants  for  yellow  and  green  given  in 
table  11  is  derived  by  dividing  the  number  of  plants  which  were  con- 
stant by  the  total  grown  of  the  pattern  concerned.  This  gives  an 
index  of  the  constancy  of  a  type,  although  it  does  not  take  into  account 
the  varying  degrees  of  the  fluctuations  which  appeared. 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS. 


43 


Judging  from  the  data  on  changes  in  green  and  yellow,  there  appears 
to  be  no  general  correlation  between  the  number  of  fluctuating  plants 
and  the  number  of  cases  of  bud  variations.  In  the  green-yellow-red 
blotched  group  there  were  proportionally  more  than  four  times  as  many 
bud  variations  as  in  the  group  green-yellow  spotted-red  blotched,  but  the 
percentages  of  constant  plants  were  nearly  identical.  Not  one  of  the 
plants  with  laciniate  leaves  was  constant  for  green  and  yellow,  but  only 
one  case  of  bud  variation  occurred.  In  the  green-red  blotched  group 
there  was  chance  only  for  the  appearance  of  yellow,  and  this  change 
occurred  in  a  relatively  large  number  of  cases,  both  in  fluctuations  and 
as  bud  variations. 

A  very  marked  contrast  appears  in  a  comparison  of  the  two  patterns 
green-yellow-red  blotched  and  yellow-green-red  blotched.  Both  have  about 
the  same  proportions  of  green  and  yellow,  except  that  the  relative 
position  is  reversed.  Both  groups  agree  quite  closely  in  the  per- 
centage of  constant  plants.  In  the  latter,  however,  bud  variations 
were  2.5  times  as  frequent.  The  position  of  the  j^ellow  in  the  center 
seemed  to  increase  bud  variations  involving  green  and  yellow  over 
that  in  plants  with  the  yellow  at  the  border. 


DISTRIBUTION  OF  BUD  VARIATIONS  AMONG  DIFFERENT  CLONES. 

The  wide  range  of  variation  both  of  fluctuations  and  of  bud  varia- 
tions emphasized  in  the  summaries  already  given  was  realized  in  a 
series  of  plants  derived  by  vegetative  propagation  from  two  plants 
having  the  same  color  pattern.  The  records  of  pedigrees  show  that 
marked  differences  appeared  among  the  various  clones  with  respect  to 
constancy  and  to  the  range  and  the  frequency  of  bud  variations. 

This  is  shown  quite  clearly  when  the  data  regarding  the  main  clones 
derived  from  plant  1  are  grouped  together  as  arranged  in  table  12. 

Table  12. — General  summary  of  clones. 


Clone. 

Total 
number 
plants. 

Plants 
constant. 

P.  ct.  of 

plants 

constant. 

Number 

of  bud 

variations. 

Ratio  of 
frequency. 

11 
12 
13 
14 
117 
111 

211 
192 
138 
155 
91 
34 

132 
87 
75 
80 
54 
29 

62 
45 
54 
51 
59 
85 

49 
21 

4 
18 
31 

4 

860 
1,830 
6,900 
1,720 

590 
1,700 

The  main  clones  11  and  12  were  derived  from  two  branches  of  plant 
1  which  had  the  same  color  pattern.  Although  the  branches  were 
identical  in  appearance,  the  two  progenies  were  quite  different.  62 
per  cent  of  clone  11  were  constant,  while  45  per  cent  of  clone  12  were 
constant;  but  in  the  more  constant  clone  11  there  were  proportionally 


44 


ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 


Table  13 

— 

Record  of  Clone  117. 

Generation 
and  numbei 
of  the  plant 

d 

'•1 

o 

o 

a 

1 

CO 

a 
o 
O 

a 

03 

3 

Bud  variations. 

Remarks. 

o 

01 

tn 

=i 
O 
« 

a 

03 

d 
o 
ft 

T3 

-d 
o 

0) 
h 
1 

".d 
d 

o 

-a 
<v 

o 

_o 

"S 

d 
<u 
<v 
u 

o 

01 

o 

t 

Hi 

1 

"3 

-6 
"o 

a) 

1 

d 

Ol 

d 
IP 

O 

-6 

O 

a 

_o 

>. 
1 

d 

Ol 

o 

13 
>> 

d 

01 

a 

1911-1912 
117 

1912-1914 
117,1 

1912-1913 
117,11.... 
117,12.... 
117,13 

117, 14. . . . 

117,15.... 

117,  16 

117,17 

1913 
117,111... 
117,112... 
117,121... 
117,122... 
117,132. .. 
117, 141. . . 
117,142... 

117,143... 

117,144... 
117,151... 
117,152... 
117,161... 
117,162... 

1913-1914 

117,111,1. 

117,111,2. 
117,  113.. . 
117,122,1. 
117, 133.. . 
117,134... 
117,135... 
117,141,1. 
117,141,2. 
117,142,1. 
117,142,2. 
117,143,1. 
117,143,2. 

117,143,3. 

117,144,1. 
117,151,  1. 
117,151,2. 

117,  151,3. 

.      2 

.      2 

2 
.      2 
.      6 

■{I 

.      6 

.      6 

6 

.      2 
2 
.      2 
.      2 
.      6 
.      2 
.      5 
r  2 

"I  5 
.      5 
.      6 
.      6 
.      6 
6 

■il 

2 
2 
2 
'      6 
.      6 
.      6 
.      2 
.      2 
.      5 
.      5 
.      2 
.      5 

r. 

.      5 

1 

.      6 

{I 

X 

X 
X 

X 

X 
X 

X 
X 
X 

X 
X 
X 

X 
X 

X 
X 
X 

X 
X 

X 
X 
X 

X 

X 
X 

X 

X 

X 
X 

X 
X 

X 

X 

X 
X 
X 

X 
X 

X 

X 

.  . 
X 

X 

X 
X 

X 

X 

X 
X 

X 

Somewhat  greener  in  winter. 

Each  part  distinct,  but  somewhat  fluctuating. 

Somewhat  greener  in  winter. 

Do. 
Mixed  during  winter,  both  greener  and  yellower. 
Constant  for  pattern,  somewhat  greener  in  winter. 
Constant  for  loss  of  yellow. 
Well-defined  sectorial  bud  variations. 
Shghtly  greener  in  winter. 

Do. 

Each  part  constant. 

Slightly  more  yellow  as  summer  advanced. 

Do. 

Do. 
Very  irregular  patterns. 
Each  part  constant. 
No  trace  of  yellow. 
SUght  fluctuation  in  yellow. 
No  trace  of  vellow. 

Do. 
Bud  variation  verj-  marked;  parts  uniform. 

Do. 

Do. 
Irregular  patterns. 

Quite  uniform  and  constant. 

No  trace  of  yellow. 

Constant  except  for  bud  variation. 

Mixed  patterns  2  and  4. 

Very  uniform  and  constant. 

Slightly  greener  in  winter. 

Do. 
Constant  except  for  bud  variation. 
Slightly  greener  in  winter. 
Much  greener  in  winter. 

No  trace  of  yellow  except  in  two  leaves  late  in  summer. 
No  trace  of  yellow  except  in  one  leaf  late  in  summer. 
Shghtly  greener  in  winter. 
No  trace  of  yellow. 
Very  constant. 

Constant  except  for  bud  variation. 
No  trace  of  yellow  except  in  two  leaves  in  late  summer. 
Died  in  a  few  weeks. 
Bud  variation  sharply  sectorial. 
Constantly  very  yellow. 
Slightly  less  yellow  in  winter. 

BY   THE    SELECTION    OF    SOMATIC    VARIATIONS. 
Table  13. — Record  of  clone  117 — Continued. 


45 


Generation 
and  number 
of  the  plants 


a 


si 


Bud  variations. 


m  \Xi 


TJ 

>) 

I 

a 

V 

(-1 

O 


tu 

^^ 

T3 

o 

I 

a> 


o  o 


Remarks. 


1913-1914 
Cont'd. 


117, 

152. 1 . 

117, 

152,2. 

117, 

152.3. 

117, 

152,5. 

117, 

152.7. 

117, 

153... 

117. 

154... 

117, 

155.  .  . 

117. 

156. .. 

117, 

157... 

117, 

161.1.  . 

117, 

161.2.. 

117, 

161,3.. 

117, 

161,4.. 

117. 

161.5.. 

117, 

162,1.. 

117, 

162,2.  . 

117, 

162,3.. 

117, 

171.... 

117, 

18 

117, 

19 

117, 

1X1... 

117, 

1X3... 

117. 

1X5... 

1914 

117, 

111,11. 

117, 

111.12. 

117, 

123.1.  . 

117, 

133, 1 .  . 

117, 

136,  1.  . 

117, 

136,2.. 

117, 

142,11. 

117, 

142,12. 

117, 

142,21. 

117, 

152,31. 

117, 

152.32. 

117, 

152,33. 

117, 

154,1.  . 

117, 

154,2.  . 

117, 

156, 1 . . 

117. 

162,11. 

/ 1 

\  6 
1 
6 
6 

114 
6 
1 
5 
6 
6 
6 
6 
6 
1 
6 
1 
6 
1 
6 
1 
6 
1 
6 
6 
6 
6 
6 
2 
5 
5 
2 
5 
2 
2 
5 

5 
2 
2 
6 
6 
6 
5 
5 
5 
14 
14 
14 
5 
6 
5 
6 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Very  yellow  at  all  times. 
Slightly  less  yellow  in  winter. 
Very  yellow  at  all  times. 
Slightly  greener  in  wdnter. 

Do. 

Do. 

Do. 
Plant  lived,  but  made  poor  growth. 
Fluctuating  to  yellow  spotted. 
Somewhat  greener  in  winter. 
Very  constant  except  for  bud  variations. 
Very  constant. 
Much  greener  in  winter. 

Do. 
Constantly  very  yellow,  but  greener  in  winter. 
Somewhat  greener  in  wanter. 

Do. 
Very  uniform  except  for  bud  variation. 
Constantly  very  yellow. 
Verj^  uniform  except  for  bud  variation. 
This  part  soon  died. 
Slightly  greener  in  winter. 

Do. 
Mixed  patterns,  4,  5.  and  6. 
Slightly  greener  in  winter. 

Do. 

Do. 

Do. 
Lived  only  a  few  weeks. 
Soon  died. 
No  trace  of  yellow. 
Very  constant. 

Verj'  constant;  no  trace  of  yellow. 
Very  constant  except  for  bud  variation. 

Fluctuated  to  yellow  spotted. 

No  trace  of  yellow. 

Slightly  more  yellow  as  summer  advanced. 

Do. 
Slightly  more  yellow,  except  for  bud  variation. 
SUght  increase  of  yellow. 

Do. 
No  trace  of  yellow  except  in  two  leaves. 
No  trace  of  yellow  except  in  one  leaf. 
No  trace  of  yellow  except  in  three  leaves  (seetext-fig.l). 
Slight  increase  of  yellow. 

Do. 

Do. 
No  trace  of  yellow  except  in  branch  with  Inid  variation. 
Slight  increase  of  yellow. 

Do. 

Do. 


46  ESTABLISHMENT    OF    VARIETIES   IN    COLEUS 

more  than  twice  as  many  bud  variations.  Clone  13,  which  was  derived 
from  a  branch  that  was  green-red  blotched,  gave  a  progeny  (of  several  pat- 
terns) of  which  54  per  cent  were  constant,  but  bud  variations  were  very 
infrequent.  Clone  14,  with  nearly  the  same  percentage  of  constant 
plants,  produced  four  times  as  many  bud  variations.  This  summary 
of  the  data  by  clones  irrespective  of  patterns  shows  a  general  irregular- 
ity and  lack  of  correlation  between  fluctuating  variations  and  bud  vari- 
ations. The  special  interest,  however,  pertains  to  the  clones  11  and  12, 
which  show  that  two  branches  apparently  identical  may  have  quite 
different  potentialities  for  constancy  and  for  bud  variations. 

Even  more  marked  differences  than  these  developed  among  the 
various  subclones.  A  study  of  pedigrees  shows  that  in  all  patterns 
and  in  all  main  clones  there  were  certain  lines  of  progeny  much  more 
constant  than  many  others.  These  could  not  be  detected  by  any  other 
than  a  pedigree  method. 

Clone  111  can  be  given  as  one  of  the  most  constant  clones.  Its 
members  numbered  34.  Four  cases  of  bud  variation  appeared;  3  were 
a  loss  of  yellow  and  1  was  a  reversal  of  the  position  of  the  green  and 
yeUow  occurring  in  one-half  of  a  leaf  only.  These  4  plants  were  other- 
wise constant.  Only  1  plant  showed  fluctuating  variability,  becoming 
quite  uniform  for  green-yellow  spotted-red  blotched.  All  the  bud  varia- 
tions involved  changes  in  the  green  and  yellow.  There  were  no  marked 
changes  in  the  amount  and  distribution  of  epidermal  red.  As  shown  in 
table  12,  the  percentage  of  constant  plants  was  85  and  the  ratio  of  bud 
variations  was  1  to  1,700.  The  clone  was  highly  constant  both  in  regard 
to  fluctuations  and  bud  variations. 

On  the  other  hand,  the  series  of  plants  derived  from  plant  117  was 
26  per  cent  lower  in  number  of  constant  plants  and  gave  nearly  three 
times  as  many  bud  variations,  yet  plants  111  and  117  were  both 
uniform  and  constant  for  the  pattern  green-yellow-red  blotched  and  were 
apparently  identical.  Until  the  autumn  of  1912,  plant  1171  was  the 
only  one  of  the  17  plants  grown  in  clone  11  that  showed  variation.  It 
gave  during  the  summer,  by  sectorial  variation  in  the  main  axis,  6 
branches  with  the  position  of  the  green  and  the  yellow  reversed.  The 
plant  was  grown  in  a  large  pot  during  the  winter  and  then  grown  out 
of  doors  during  the  following  summer.  In  the  second  sunmier  two 
more  bud  variations  appeared  on  the  part  with  green-yellow-red  blotched 
foliage,  but  on  branches  quite  separated.  Both  were  sectorial;  one  was  a 
loss  of  green,  giving  the  yellow-red  blotched  pattern,  and  one  was  a  loss 
of  yellow,  giving  the  green-red  blotched  pattern.  The  plant  possessed  for 
some  time  four  patterns,  each  uniform  for  a  certain  part  of  the  plant. 

The  record  of  pedigree  for  the  progeny  of  the  plant  can  be  given  as 
illustrating  a  clone  in  which  bud  variations  occurred  with  a  high  ratio 
of  frequency.  In  table  13  the  plants  are  arranged  in  generations 
according  to  number.     To  trace  the  progeny  or  the  ancestry  of  any 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  47 

plant,  one  should  look  in  the  generations  following  or  preceding  for  the 
serial  numbers.  Certain  plants  were  grown  with  two  patterns  as  a 
chimera,  and  these  are  indicated  by  brackets,  with  a  record  for  each 
pattern.  In  order  to  make  the  tabulation  more  compact,  numbers 
are  used  to  represent  the  different  patterns,  and  these  correspond  with 
the  numbers  of  the  figures  in  the  plate,  as  follows: 

1  =  yellow-red  blotched.  5  =  green-red  blotched. 

2  =  green-yellow-red  blotched.  6  =  yellow-green-red  blotched. 
4  =  green-yellow  spotted-red  blotched.      14  =  yellow-green. 

A  survey  of  this  series  of  plants  shoAvs  that  on  the  91  plants  31  bud 
variations  appeared,  giving  a  ratio  of  frequency  of  1  to  590  against 
1  to  860  for  the  entire  clone  11  and  1  to  1,700  for  the  sister  clone  111. 

A  further  analysis  within  this  progeny  shows  that  in  several  cases 
similar  bud  variations  can  be  traced  to  a  common  ancestry.  For 
example,  6  of  the  9  cases  of  what  has  been  called  spontaneous  develop- 
ment of  3^ellow  occurred  during  the  summer  of  1914  in  plants  descended 
from  plant  117142  (see  table  6),  which  itself  was  constant  for  loss 
of  yellow  during  its  period  of  growth.  Another  case  of  spontaneous 
development  of  yellow  was  in  plant  1171441.  In  this  clone,  therefore, 
all  cases  of  spontaneous  development  of  yellow  were  in  plants  descended 
from  plant  11714. 

It  is  quite  clear  from  such  pedigrees  that  distinct  differences  in 
tendencies  in  regard  to  the  degree  of  variation  may  exist  among  buds 
of  branches  bearing  similar  foliage. 

ENVIRONMENTAL  INFLUENCE. 

Observations  were  made  and  pedigrees  of  plants  examined  to  deter- 
mine whether  changes  in  ordinary  environmental  conditions  influence 
fluctuations  and  bud  variations. 

To  secure  accurate  data  on  the  relative  frequency  of  bud  variations 
during  summer  and  winter  is  hardly  feasible.  In  general  only  about 
half  of  the  summer  plants  were  from  cuttings  made  early  in  the  spring. 
The  others  were  grow^n  in  pots  in  a  greenhouse  during  the  winter  and 
transplanted  to  the  garden,  where  the}'  grew  during  the  summer, 
making  larger  plants  with  many  more  branches  than  were  produced 
during  the  winter.  Of  the  total  number  of  1 15  bud  variations  iuA'olving 
color,  9  appeared  during  the  winter,  which  is  fairly  proportional  to  the 
relative  number  of  branches  that  developed. 

There  was  a  strong  tendency'  for  plants  having  yellow  to  become 
greener  in  winter  and  yellower  in  summer,  and  also  to  become  greener 
when  severely  pruned.  At  any  time,  however,  during  the  winter, 
some  plants  of  each  pattern  having  yellow  (for  example  green-yellow- 
red  blotched)  could  be  found  with  the  pattern  as  pronounced  as  dur- 
ing the  summer.  During  the  winter  of  1913-14,  two  plants  of  each  of 
the  following  types,  green-yellow-red  blotched,  green-yellow  spotted-red 


48  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

blotched,  and  green-yellow-solid  red,  were  grown  in  a  greenhouse  with 
northern  exposure  only,  which  gave  scarcely  any  direct  lighting.  These 
plants  were  constant  for  the  respective  patterns.  The  green  and 
yellow,  however,  were  slightly  less  bright  and  intense. 

During  the  summer  of  1914  several  plants  of  nearly  all  types  of 
pattern  containing  yellow  were  grown  in  a  greenhouse  the  glass  of 
which  was  whitewashed  to  decrease  the  intensity  of  illumination.  No 
appreciable  differences  could  be  noticed  between  the  patterns  of  these 
and  of  plants  grown  out  of  doors.  So  far  as  I  have  observed,  it  does 
not  seem  that  any  of  the  color  variations  can  be  attributed  to  such  fac- 
tors as  heat  or  degree  of  illumination. 

Furthermore,  the  loss  of  yellow,  loss  of  green,  and  gain  and  loss  of 
red  all  occurred  in  single  branches  and  in  sections  of  branches  (figs. 
21  and  24).  Frequently  two  quite  difTerent  changes  appeared  on  the 
same  plant,  which  was  then  grown  for  some  time  with  2,  3,  or  even 
4  quite  distinct  types  of  foliage.  Cuttings  were  made  so  as  to  give 
plants  with  two  types  of  foliage,  as  (a)  green  and  green-red  blotched,  or 
(6)  green-yellow-red  blotched  and  green-red  blotched,  or  (c)  green-yellow-red 
blotched  and  green-yellow-solid  red,  etc.  In  all  these  cases  branches 
with  two  types  of  foliage  were  submitted  to  as  uniform  conditions 
as  possible;  they  grew  on  the  same  plant,  were  subject  to  the  same 
degree  of  heat  and  illumination,  and  were  supphed  by  the  same  root 
system.  Under  this  test  the  different  patterns  were  fully  as  constant 
as  if  grown  on  separate  plants. 

These  facts  indicate  quite  clearly  that  the  marked  and  sudden 
variations  and  differences  in  expression  of  color  concerned  in  the 
different  patterns  are  not  readily  attributable  to  external  environ- 
mental factors. 

Flammarion  (1898)  used  a  variety  of  Coleus  with  a  yellow,  green, 
and  red  color  pattern  in  testing  the  influence  of  hght  on  pigmentation 
in  plants.  With  red  light  he  secured  decrease  of  red  pigmentation 
and  a  broader  leaf;  under  influence  of  green  light  the  red  coloration 
mostly  disappeared,  and  under  blue  light  there  was  somewhat  less  red. 
A  series  grown  out  of  doors  under  conditions  of  diffused  hght  showed 
decrease  of  red  coloring,  while  those  under  very  dim  light  gave  still 
less  development  of  it.  With  decrease  of  red  the  center  of  the  leaves 
became  quite  yellow.  Evidently  the  red  pigmentation  of  his  variety 
was  chiefly  located  in  the  epidermis.  In  marked  contrast  to  these 
results  it  may  be  noted  that  the  bud  variations  that  I  have  reported 
give  more  marked  changes  than  those  induced  by  Flammarion  and 
that  these  appear  suddenly  and  in  a  sector  of  a  bud  in  a  manner  that 
suggests  internal  readjustments  rather  than  external  environmental 
influence. 

In  respect  to  the  laciniate  leaf-shape  and  its  periodic  development, 
however,   environmental  influences  seem  to  have  some  effect.     As 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  49 

already  indicated,  conditions  favoring  rapid  and  vigorous  growth  lead 
to  the  development  of  entire  leaves.  In  vegetative  propagation, 
however,  the  periodic  laciniate  condition  develops  only  in  certain 
subclones.  In  other  subclones  it  has  not  appeared.  That  is,  the  same 
conditions  of  environment  and  treatment  do  not  lead  to  the  appear- 
ance of  the  laciniate  character  in  all  plants,  so  it  is  hardly  to  be  con- 
sidered as  purely  environmental.  The  laciniate  character  has  also 
developed  in  one  plant  in  the  manner  of  bud  variations. 

SEED  PROGENY. 

The  data  obtained  from  the  seed  progeny  of  my  strains  of  Coleus 
have  direct  bearing  on  the  nature  and  inheritance  of  the  bud  variations 
that  appear  and  indicate  that  bud  variations  can  give  rise  to  as  widely 
difTerent  forms  as  can  be  obtained  among  the  various  members  of  a 
hj^brid  progeny. 

Selfed  seed  was  obtained  from  a  plant  of  the  pattern  yellow-green- 
red  blotched  and  22  plants  were  grown  during  the  latter  part  of  the 
summer  of  1914,  In  respect  to  the  development  of  green  and  yellow, 
there  was  every  gradation  between  green  with  large  yellow  blotches 
irregularly  distributed  through  the  leaf-blade  and  pure  green.  In 
respect  to  the  development  of  red  in  the  epidermis,  there  were  grada- 
tions from  absence  of  red  to  a  general  distribution  of  large  irregular 
blotches.  As  in  the  case  of  bud  variations,  the  difTerent  types  of  epi- 
dermal red  occurred  independently  of  the  degree  of  development  of 
underlying  green  and  yellow. 

In  regard  to  leaf-shape,  there  was  every  range  of  variation.  Few 
plants  could  be  classed  as  laciniate,  but  there  was  every  gradation  from 
shallow  to  deep  lobing  and  from  coarse  to  fine  lobing.  The  leaves  on 
any  one  plant  were  quite  uniform.  It  should  be  noted  that  the  lacini- 
ate character  had  not  appeared  in  the  particular  subclone  from  which 
these  seedlings  were  derived.  It  had  appeared  in  sister  subclones 
as  described  above.  Seven  plants  possessed  leaves  quite  like  those  of 
the  parent  type,  both  in  respect  to  the  cuneate  base  and  the  crenate 
margin.  Five  plants,  however,  had  large  leaves,  some  measuring  10 
inches  long,  that  w^ere  broadly  obtuse  at  the  base,  with  the  cuneate 
character  lacking.  From  plants  possessing  fiat  leaves  with  a  smooth 
surface  there  was  gradation  to  those  with  leaves  much  crinkled  or 
folded. 

At  the  same  time  23  plants  were  grown  from  selfed  seed  of  a  plant 
which  possessed  the  laciniate  leaf-type  as  a  fluctuating  character. 
The  plant  itself  also  fluctuated  in  respect  to  the  development  of  green 
and  yellow  as  follows:  during  the  winter  of  1913-14,  its  leaves  were 
strongly  laciniate  and  devoid  of  yellow,  and  during  the  following  sum- 
mer the  plant  was  quite  yellow,  becoming  almost  like  the  type  green- 
yellow-red  blotched  (fig.  2)  and  every  leaf  was  entire.     In  their  develop- 


50  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

ment  of  green  and  yellow  the  plants  ranged  from  pure  yellow  plants 
that  died  within  a  few  weeks  to  those  that  were  pure  green.  Two 
of  the  plants  had  much  yellow  in  very  irregular  and  mixed  patterns. 
Eleven  had  no  trace  of  yellow  for  several  months,  when  a  few  yellow 
spots  appeared  in  some  of  the  leaves  on  several  plants.  In  regard  to 
epidermal  red  there  were  numerous  types  ranging  from  solid  red  to  no 
red.  Of  those  wdth  red-blotched  epidermis,  some  were  uniformly 
finely  blotched,  while  on  others  the  blotches  were  large,  with  a  single 
blotch  sometimes  covering  one-fourth  of  a  leaf.  As  to  the  shape  of  the 
leaves,  the  series  showed  the  same  range  of  variation  exhibited  by  the 
seed  progeny  described  above.  There  were  but  5  that  were  strongly 
laciniate. 

Besides  the  above  plants  which  were  grown  to  maturity,  there  are 
at  the  present  date  (February  8,  1915)  20  seedlings  of  a  plant  that 
had  an  epidermis  of  solid  red,  as  shown  in  figure  8.  The  plants  have 
from  2  to  3  pairs  of  leaves,  but  it  is  clear  that  only  one  of  the  seedhngs 
has  a  solid  red  epidermis.  Red-blotched  types  prevail  and  few  of  the 
seedlings  show  any  yellow  coloration. 

Summarizing,  it  is  plain  that  the  plants  grown  from  seed  give  wide 
variations.  In  respect  to  color  patterns,  there  were  numerous  types 
which  gave  very  complete  gradations  between  extremes,  especially  in 
regard  to  epidermal  red.  Many  of  the  types  that  had  appeared  as 
bud  variations  appeared  also  in  the  seed  progenies,  such  as  yellow-red 
blotched,  green-yellow  spotted-red  blotched,  green-red  blotched  (wide  range 
of  variation  in  respect  to  size  of  blotches),  green-yellow  spotted,  and  green. 
One  of  the  new  patterns  could  be  described  as  yellow-green  blotched- 
solid  red.  Another  had  the  red  blotches  of  the  epidermis  coalescing  at 
certain  points,  making  the  red  markings  continuous  but  not  solid  (see 
fig.  29),  so  that  the  underlying  green  showed  through  in  blotches. 

In  vegetative  propagation  leaf-shape  remained  very  constant,  the 
only  exception  being  the  clones  that  developed  laciniate  leaves.  In 
the  seed  progeny  of  all  plants  tested,  however,  new  types  of  entire 
leaves  have  appeared.  The  laciniate  leaf  characters  and  various 
intermediates  between  it  and  the  typical  entire  leaf  appeared  in  the 
first  leaves  of  certain  plants  and  remained  as  a  rather  constant  character 
during  the  time  the  plants  have  been  grown. 

The  wide  variations  appearing  in  the  seed  progeny  indicate  that  this 
strain  of  Coleus  is  either  of  mixed  parentage  or  that  the  processes 
concerned  with  production  of  color  patterns  and  leaf-shape  are  them- 
selves subject  to  wide  variations.  The  variations  in  respect  to  color 
patterns  were,  however,  no  greater  in  range  or  extent  than  were  those 
that  appeared  in  bud  variations,  and  the  fluctuations  from  entire 
to  laciniate  leaf-shape  gave  extreme  types  of  leaf-shape,  with  all  grades 
of  intermediates,  on  a  single  plant,  as  is  quite  well  shown  in  plant  123153 
of  plate  4. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  51 

HISTORY  OF  COLEUS. 

The  early  historj^  of  the  cultivated  varieties  of  Coleus  shows  that  the 
original  species  utilized  in  cultivation  and  hj'bridization  were  few  in 
number  and  relatively  simple  in  respect  to  diversity  of  color  patterns 
and  leaf-shape.  The  variations  which  appeared  in  vegetative  and  in 
seed  propagation  within  a  few  years  after  introduction  gave  a  wide 
range  of  variabiUty,  with  greater  extent  than  the  extremes  of  the  char- 
acters in  the  original  species.  In  general  the  variations  reported  in  this 
paper  are  quite  parallel  with  those  appearing  in  the  development  of  the 
numerous  cultivated  varieties,  both  in  regard  to  the  apparently  spon- 
taneous development  of  new  patterns  and  to  the  reversion  to  parent 
or  ancestral  types. 

It  appears  that  the  first  variegated  species  of  Coleus  introduced 
into  European  cultivation  was  Coleus  blumei  Bentham.  The  original 
description  (Blume,  1826)  under  the  name  Plectranthus  scutellarioides 
states  that  the  leaves  were  spotted  above  with  dark  purple  (folio 
supra  maculis  atropurpureis  picta).  This  plant  was  introduced  into 
Holland  in  1851  and  the  next  year  Planchon  (1852)  gave  a  brief  de- 
scription of  it,  accompanied  by  a  colored  plate.  It  was  soon  intro- 
duced into  England,  and  in  1853  a  description  with  colored  plate 
appeared  in  an  English  magazine  (Hooker,  1853).  These  two  illustra- 
tions agree  quite  closely,  although  the  latter  shows  the  plant  in  some- 
what a  brighter  green,  with  leaves  with  a  somewhat  more  solid  mass  of 
central  red.  The  central  part  of  the  upper  surfaces  were  dark  purple 
or  sanguineous,  breaking  into  spots  near  the  margin.  In  the  descrip- 
tion it  is  stated  that  the  leaves  were  entire  at  the  base,  ''which  is  gradu- 
ally attenuated  into  a  more  or  less  elongated  petiole,"  a  character  well 
shown  in  an  outUne  drawing  of  a  leaf. 

There  are  no  specimens  of  this  species  in  the  Bentham  collections 
at  the  Kew  Herbarium.  There  is,  however,  a  specimen  in  Herb. 
Hookerianum  with  a  label  in  Sir  WiUiam  Hooker's  handwriting,  stating 
that  the  plant  w^as  grown  at  Kew  Gardens  and  citing  the  description 
and  plate  in  Botanical  Magazine  (Hooker,  1853).  This  was  evidently 
regarded  by  Hooker  as  a  typical  specimen.  The  leaf-shape  is  identical 
with  that  of  the  illustration  referred  to  and  is,  with  the  exception  of  fig- 
ure 7,  quite  the  same  as  the  leaves  shown  in  the  plates  illustrating  this 
paper.  From  the  description  and  illustration  the  color  pattern  was 
nearly  identical  with  my  type  green-upper  center  solid  red  (fig.   15). 

Morren  (1856)  describes  the  variety  C.  blumei  pectinatus  as  somewhat 
more  richly  colored,  but  differing  chiefly  in  having  the  leaves  deeply 
and  doubly  lobed.  The  colored  plate  shows  that  the  base  of  the 
leaf-blade  was  cuneate,  as  in  the  species. 

Coleus  verschaffeltii  was  first  named  by  Lemoine  (1861),  who  decided 
that  it  was  distinct  from  C.  blumei.  It  appears  (Witte,  1862)  that  this 
plant  was  introduced  into  Rotterdam  in  1860  from  Java.     Colored 


52  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

plates  (111.  Hort.,  8,  pi.  293,  and  Flor.  Mag.,  2,  pi.  96)  show  that  this 
species  was  richly  and  deeply  colored  with  crimson  on  both  surfaces 
and  that  the  base  of  the  leaf -blades  were  not  cuneate  but  heart-shaped. 
Being  richer  in  color  than  C.  hlumei,  this  plant  attracted  considerable 
attention  as  a  foliage  plant.  In  1864  (Gard.  Chron.,  p.  506),  a  sport 
of  this  species  called  marmoratus  with  bright  green  patches  in  its  leaves, 
was  described. 

Concerning  C.  verschaffeltii,  Herincq  (1865)  remarks  that  the  expres- 
sion of  red  coloration  fluctuates  with  light  conditions  and  suggests 
that  one  might  eliminate  the  red  by  keeping  the  plants  in  the  shade. 
Later  he  (1866)  notes  fluctuations  that  are  less  due  to  environment, 
for  he  observes  variation  in  the  leaves  of  a  single  plant  and  states  that 
no  doubt  selection  of  cuttings  would  give  pure-green  plants.  He  men- 
tions that  he  had  seen  a  young  plant  having  no  trace  of  red  coloration. 

Two  types  of  Coleus  destined  to  play  an  important  part  in  the 
development  of  horticultural  varieties  were  introduced  into  England 
from  New  Caledonia  by  John  G.  Veitch.  Although  briefly  mentioned 
in  1866,  they  were  first  described  and  illustrated  in  1867.  C.  gibso7iii 
(Verlot,  1866 ;  Dombrain,  1867a)  was  of  a  dwarf  bush}^  habit.  The  leaves 
were  large  and  ''of  a  light-green  color,  distinctly  veined  and  blotched 
with  dark  crimson-purple."  The  plate  clearly  shows  that  the  leaves 
were  only  slightly  crenate  and  that  the  bases  were  broadly  cordate. 
C.  veitchii  (Dombrain,  18676)  possessed  leaves  quite  similar  in  shape, 
but  with  the  entire  central  portion  of  the  leaf  of  a  deep  chocolate  color 
with  the  edges  green. 

These  four  species,  C.  hlumei,  C.  verschaffeltii,  C.  gibsonii,  and  C. 
veitchii,  were  used  as  parents  in  the  production  of  hybrids  by  F.  Bause, 
in  the  employ  of  the  Royal  Horticultural  Society  of  London.  12 
hybrids,  of  which  C.  verschaffeltii  was  the  seed  parent,  resulted  the  first 
year.  Rather  extended  descriptions  of  these  are  given  by  Thomas 
Moore  (1868).  The  Fi  progeny,  even  from  the  same  parentage,  were 
widely  different,  some  resembling  the  seed  parent,  while  others  resem- 
bled the  pollen  parent.  In  regard  to  leaf-shape,  there  were  two  groups, 
one  with  flat  crenate  leaves,  as  C.  veitchii,  and  one  with  frilled-dentate 
leaves,  as  C.  verschaffeltii.  In  colors  there  were  various  shades  of  purple 
in  solid  colors,  blotched  areas,  and  in  reticulations.  At  the  time  this 
variation  among  the  Fi  progeny  aroused  considerable  interest.  One 
anonymous  writer  (Gard.  Chron.,  33,  407)  raises  the  question  how 
several  kinds  of  Coleus  could  originate  from  the  same  cross.  But 
one  hybrid  with  C.  hlumei  as  a  parent  is  reported.  This  had  frilled 
leaves  and  coloration  much  like  that  of  C.  hlumei.  It  was  less  deeply 
colored  than  the  hybrids  resulting  for  the  other  pollen  parents. 

There  is  no  mention  of  yellow  in  any  of  these  hybrids.  All  were 
bi-colored,  but  with  striking  combinations  of  the  green  and  various 
shades  and  amounts  of  purplish  or  red  colorations.     They  were  sold 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  53 

at  auction  (Gard.  Chron.,  33,  432)  for  the  aggregate  sum  of  £390. 
One  hybrid  brought  59  guineas. 

The  production  of  these  valuable  variegated  Coleus  varieties  stimu- 
lated further  hybridization  work.  Wilham  Bull  produced  18  different 
types  (reported  by  Herincq,  1868),  and  in  November  of  that  year  he 
advertised  pedigreed  seed  from  20  crosses  involving  14  varieties  grown 
that  year  (Advertisement,  Gard.  Chron.,  33,  1232). 

Meanwhile,  at  the  gardens  of  the  Royal  Horticultural  Society,  a 
new  series  of  hybrids  were  produced  much  finer  than  those  of  the 
previous  year.  The  parentage  of  these  interesting  hybrids  is  not  fully 
given,  but  it  is  stated  by  Moore  (1869)  that  certain  of  the  hybrids  of 
the  previous  year  were  crossed  with  C.  hlumei  itself.  In  this  second 
lot  of  hybrids,  yellow  coloration  appeared  as  a  new  or  spontaneous 
development.  Eight  (Gard.  Chron.,  33,  1210)  possessed  distinct 
yellow,  forming  in  some  cases  a  golden  margin.  Two  (Prince  Arthur 
and  Princess  Beatrice)  are  described  as  having  a  yellowish  ground- 
color or  golden  green.  Most  of  them  had  a  yellow  and  green  ground- 
color overlaid  with  shades  of  purple  or  crimson  red.  The  most  brilliant 
of  the  series  was  named  Queen  Victoria,  a  colored  plate  of  which 
appeared  as  a  frontispiece  in  the  Florist  and  Pomologist  (volume  for 
1869).  This  plate  shows  that  the  ground-color  was  mostly  yellow, 
overlaid  by  an  epidermal  red,  appearing  crimson  over  the  yellow  and 
entirely  covering  the  upper  surface  of  the  leaves  except  at  the  margin. 
None  of  the  series  possessed  frilled  leaves.  The  leaves  of  all  w^ere 
flat,  with  crenate  teeth  somewhat  deeply  cut. 

Although  of  much  more  remarkable  variegation  than  the  hybrids  of 
the  previous  year,  9  of  these  new  coleuses  brought  but  65  guineas. 

While  the  yellow  element  in  the  variegation  appeared  strongly  in  this 
second  lot  of  hybrids,  it  should  be  noted  that  in  the  year  1867  (Gard. 
Chron.,  33,  460)  a  golden  Coleus  arose  as  a  bud-sport  from  C.  hlumei. 
It  is  described  as  like  C.  hlumei,  but  with  the  green  exchanged  for  a 
decided  yellow  tint.  The  sport  appeared  in  one-half  of  a  single  leaf. 
The  bud  at  its  base  was  propagated  and  gave  the  new  variet3^  It 
does  not  appear  that  this  sport  was  used  in  the  hybridization  work  that 
produced  the  golden  coleuses. 

During  1868  and  1869,  the  various  horticultural  publications  men- 
tion by  name  no  less  than  54  new  varieties  of  Coleus.  For  several 
years  thereafter  few  varieties  were  mentioned,  but  in  1878  (Garten- 
flora,  p.  50)  13  forms  not  previously  mentioned  are  listed.  The  next 
year  this  journal  (pp.  341-346)  states  that  breeding  of  Coleus  had  been 
carried  on  in  Germany,  speaks  of  new  forms  that  arose,  and  prints  an 
uncolored  plate  illustrating  4  types.  New  types  were  also  credited  to 
Bull  (Rev.  Hort.  Belg.,  5:  49;  Gard.  Chron.,  45:  748). 

At  the  exposition  in  Paris  in  1879,  Morlet  exhibited  varieties  described 
by  Andre  (1879)  as  far  surpassing  all  previous  varieties.     These  had 


54  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

enormous  foliage,  with  remarkable  combinations  of  shades  of  carmine, 
yellow,  and  green.  20  types  are  mentioned  by  name  and  12  of  these 
are  described. 

Pynaert  (1881)  states  that  about  250  varieties  of  Coleus  had  been 
put  upon  the  market.  He  notes  that  it  is  difficult  to  establish  a  nomen- 
clature for  the  various  Coleus  varieties.  He  describes  and  illustrates 
in  color  a  variety  called  Reine  des  Beiges  which  arose  from  seed  of  the 
variety  Duchess  of  Edinburgh,  a  variety  illustrated  in  the  Floral 
Magazine  in  1874.  The  leaves  of  the  Duchess  of  Edinburgh  pos- 
sessed a  yellow  border;  the  Reine  des  Beiges  had  the  yellow  in  the 
center  of  the  leaves.  The  relative  position  of  the  green  and  the 
yellow  was  therefore  reversed,  which  is  the  difference  between  tj^pes 
green-yelloiv-red  blotched  and  yellow-green-red  blotched  already  reported 
in  this  paper. 

From  the  evidence  at  hand  it  is  clear  that  a  large  number  of  types  of 
Coleus  have  been  produced.  Probably  the  same  type  or  nearly  identical 
types  have  been  given  different  trade  names.  With  the  exception  of 
the  first  hybrids  produced  by  Bause,  there  is  almost  no  record  of  the 
parentage  of  most  types.  The  plants  attracted  attention  solely  on 
account  of  their  variegated  foliage,  and  for  a  time  were  more  extensively 
used  as  bedding  and  foliage  plants  than  they  are  at  the  present  time. 

From  the  standpoint  of  genetics,  it  is  suggestive  that  such  wide 
variation  appeared  in  the  cultivation  and  hybridization  of  the  4  species 
already  discussed,  although  it  should  be  noted  that  there  is  the  possibil- 
ity that  other  species  were  concerned  in  the  parentage  of  some  of  the 
varieties  now  in  cultivation.  The  parent  species  possessed  a  green 
background,  or  at  least  were  without  pronounced  yellow.  The  epider- 
mis especially  was  more  or  less  colored  with  purple  or  red  in  blumei, 
verschaffeltii,  and  veitchii,  while  in  gibsonii  the  purphsh  coloring  was 
largely  confined  to  the  veins. 

In  the  varieties  derived  from  these,  yellow  appeared  as  a  pronounced 
part  of  the  coloration.  Some  tjqjes  were  largely  yellow,  others  were 
pale  yellow,  and  others  were  entirely  green.  In  many  the  yellow  was 
localized  at  the  border,  but  in  others  it  was  at  the  center  and  in  others 
the  yellow  blotches  were  well  distributed.  These  variations  in  yellow 
and  green  were  combined  with  variations  in  amount  and  quality  of 
epidermal  and  internal  (chiefly  in  veins)  development  of  purplish  and 
red  tints. 

The  historical  evidence  indicates  that  the  form  of  Coleus  used 
in  the  experiments  here  reported  is  derived  from  C.  blumei.  In 
respect  to  the  cuneate  base  of  the  leaf-blades  and  the  marginal  charac- 
ter of  the  first  pair  of  lateral  veins  almost  making  the  petiole  attenuate, 
the  leaves  of  all  types  but  the  laciniate  agree  almost  exactly  with 
that  of  the  figure  for  C.  blumei  (Bot.  Mag.,  1853).  This  character  dif- 
ferentiated sharply  C.  blumei  from  C.  verschaffeltii,  C.  veitchii,  and  C. 
gibsonii  or  any  other  species  introduced  into  cultivation,  and  seems  to 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  55 

indicate  clearly  that  the  type  in  question  is  derived  more  or  less 
directly  from  C.  hlumei.  The  flowers  agree  with  the  description  of 
C.  hlumei. 

Coleus  hlumei,  as  stated  above,  was  already  in  cultivation  in  Java 
when  described  by  Blume.  Blume  (1826)  suggests  that  the  plant 
named  by  him  Plectranthus  laciniatus  may  have  been  simply  a  variety, 
as  it  seems  to  have  differed  largely  in  having  laciniate  leaves.  Blume's 
original  description  speaks  of  the  leaves  of  C.  hlumei  as  spotted  above, 
but  the  colored  plates  appearing  in  1852  and  1853  show  the  greater 
part  of  the  upper  surface  of  a  soUd  purphsh  color.  Coleus  hlumei  was 
introduced  into  the  German  gardens  under  the  name  Plectranthus 
concolor  var.  picta  (Gartenflora,  1853,  2:  220).  Only  in  one  of  Bause's 
first  lot  of  hybrids  was  C.  hlumei  concerned,  but  this  species  was,  it  is 
stated,  the  seed  parent  of  the  second  lot  of  hybrids.  If  this  hybrid  was 
used  in  further  hybridization  work  there  is  no  record.  It  was  the  least 
brilliantl}'  colored,  possessed  no  yellow,  and  sold  for  the  sum  of  5  guineas, 
which  was  the  lowest  sum  paid  for  any  one  of  the  12  hybrids  (Gard. 
Chron.,  3>o,  432).  I  have  been  unable  to  find  further  mention  of  this 
hybrid,  which  was  named  reevesii. 

C.  hlumei  produced  in  1868  a  bud  sport  with  the  green  changed  to  a 
decided  yellow  tint.  Through  propagation  this  gave  rise  to  the  variety 
telfordi  aurea  (Gard.  Chron.,  33,  460). 

Andre  (1880)  illustrates  and  describes  4  new  varieties,  which  he 
attributes  to  C.  hlumei.  Apparently  all  have,  however,  leaves  with 
cordate  bases  and  not  at  all  cuneate,  as  is  the  case  in  C.  hlumei,  which 
makes  his  determination  of  doubtful  validity.  The  same  is  true  of  the 
type  Gloire  de  Dijon,  described  and  figured  by  Rodigas  (1888).  Rodi- 
gas  later  (1892)  notes  the  wide  variability  obtained  from  the  seed  prog- 
eny of  what  was  considered  as  C.  hlumei  from  Chile.  The  4  derived 
types  illustrated  possess,  however,  strongly  cordate  leaves  which  make 
the  identity  with  C.  hlumei  doubtful. 

At  the  present  time  it  does  not  appear  that  any  pure  strains  of  C. 
hlumei,  C.  gihsonii,  or  C.  veitchii  are  in  cultivation.  The  strain  used 
in  these  experiments  agrees  most  closely  in  regard  to  leaf-shape  with 
the  original  C.  hlumei,  but  the  variability  of  the  seed  progeny  seems 
to  indicate  that  it  is  not  a  pure  strain.  The  writer  inquired  about  and 
observed  all  types  of  Coleus  available  at  numerous  botanical  gardens 
and  nurseries  during  a  6  weeks'  trip  to  Germany,  Holland,  and  Eng- 
land during  the  summer  of  1914.  Only  one  plant,  a  plant  observed 
at  the  Royal  Botanic  Gardens  at  Regent's  Park,  London,  was  seen 
which  had  the  hlumei  type  of  leaf. 

Coleus  verschaffeliii  is,  however,  quite  generally  in  cultivation  at  the 
present  time  and  agrees  quite  closely  with  the  type  first  described. 

I  am  especially  indebted  to  Mr.  F.  J.  Chittenden,  of  the  Royal 
Horticultural  Society,  for  securing  a  statement  (a  letter  to  Mr.  Chit- 
tenden) from  Mr.  B.  Wynne,  in  which  he  states  that  he  spent  3  months 


56  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

of  his  time  (as  a  student)  at  Chiswick  in  1866,  under  Bause,  in  the 
propagating  department.  He  was  quite  famihar  with  the  methods  used 
in  the  development  of  the  hybrid  coleuses  and  states  that  "it  fell  to  my 
lot  to  convey  the  first  half  dozen  coleuses  sold  to  Sleven's  rooms." 
Mr.  Wynne  refers  to  the  description  (see  reference  above)  of  the  hybrids 
as  adequate.  In  regard  to  the  present  existence  of  these  hybrid  types, 
Mr.  Wynne  gives  the  following  statement: 

"I  am  unable  to  say  whether  there  is  more  than  one  of  the  Chiswick-raised 
set  of  Coleus  in  existence  now,  but  I  very  much  doubt  it.  It  is  very  interest- 
ing, however,  that  at  least  one  of  them  has  survived  and  is  still  grown  for 
Covent  Garden  and  possibly  for  other  markets.  This  is  the  variety  originally 
named  Queen  Victoria,  which  received  a  first-class  certificate  (R.  H.  S.)  in  the 
autumn  of  1868,  and  was  bought  at  the  second  sale  by  the  now  extinct  firm  of 
John  and  Charles  Lee,  of  Hammersmith.  I  do  not  think  it  is  now  known  in 
the  market  by  its  original  name,  but  it  is  the  well-known  variety  with  choco- 
late leaves  and  golden  segments  and  I  have  no  doubt  about  its  identity  with 
our  old  Chiswick  plant." 

It  should  be  noted  that  Wj^nne's  description  of  this  form  as  "choco- 
late leaves  with  golden  segments"  does  not  agree  with  the  colored 
plate  in  the  Florist  and  Pomologist  for  1869.  The  latter  represents  the 
pattern  as  solid  crimson  in  the  center,  with  well-defined  yellow  border. 

While  Coleus  is  now  regarded  with  less  favor  than  it  formerly 
received,  there  are  many  types  in  cultivation  exhibiting  great  range 
of  color  patterns  and  leaf  character.  At  Erfurt,  Germany,  during 
the  summer  of  1914,  the  writer  saw  large  collections  of  Coleus  grown  for 
seed  for  the  trade.  In  the  Ernst  Benary  collections  they  were  chiefly 
of  large-leaved,  small-leaved,  and  f ringed-leaved  tj^pes.  In  the  large- 
leaved  plants  the  yellow,  if  present,  was  usually  in  the  center,  a  condi- 
tion which  was  true  without  exception  for  the  plants  with  small  leaves. 
The  fringed-leaved  types  possessed  most  curious  doubly  cut  prolifera- 
tions about  the  margin.  Of  the  entire  collection  hardly  any  two  plants 
were  ahke  as  to  color  distribution.  There  were,  however,  fewer  classes 
in  regard  to  leaf-shape.  One  of  the  newer  tj-pes  was  a  dwarf  with 
fringed  leaves  derived  by  selection. 

At  the  greenhouses  of  Haage  and  Schmidt  there  were  2  rather  definite 
types  with  laciniate  leaves.  One  is  salicifolia,  with  narrow,  slender, 
quite  irregularly-lobed  leaves.  Another  is  quercifolia,  with  broad 
leaves  coarsely  cut  with  round-tipped  segments.  Their  large-leaved 
types,  entire  and  fringed,  showed  great  diversity  of  color  patterns. 
The  plants  are  grown  for  seed,  and  in  producing  stock  for  seed  parents 
selections  are  made  with  special  reference  to  leaf-shape  and  general 
habit  of  growth.  In  general  no  selection  is  made  with  reference  to  color 
patterns  and  each  type  exhibits  wide  variation  in  this  respect. 

From  the  history  of  Coleus  it  seems  quite  clear  that  the  numerous 
and  diverse  varieties  have  arisen  from  few  species.  These  varieties 
exhibit  many  characteristics  of  coloration  and  leaf-shape  that  were 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  57 

not  possessed  by  any  of  the  parents;  for  example,  there  is  no  evidence 
that  a  yellow  element  of  variegation  was  present  in  any  of  the  parent 
species.  In  leaf-shape  also,  many  new  types  have  arisen.  Hybridi- 
zation and  rather  intense  and  artificial  cultivation  have  been  associated 
with  the  development  of  such  diversity  from  plants  comparatively 
simple. 

Considering  the  various  types  of  Coleus  as  a  whole,  we  may  note 
that  the  wide  range  of  variability  which  they  exhibit  is  in  large  measure 
realized  in  the  bud  variations  that  have  appeared  from  the  single 
types  here  reported.  The  green,  green-yellow,  yellow-red  blotched, 
and  green-solid  red  are  extreme  forms  that  are  in  degree  and  quality 
counterparts  of  the  extremes  seen  in  the  different  varieties.  In  the 
development  of  the  laciniate  character  of  the  leaf-shape  in  which  the 
leaves  on  a  single  plant  fluctuate  from  entire  to  extremely  laciniate, 
the  counterparts  of  cut-leaved  quercifolia  and  salicifolia  types  are 
in  large  degree  realized. 

In  seed  progeny  and  in  bud  variations  the  single  strain  of  Coleus 
investigated  has  in  the  3  years  of  observation  shown  the  same  types  of 
variation  that  have  developed  in  the  entire  series  of  cultivated  varie- 
ties derived  by  both  seminal  and  vegetative  variations. 


58  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

DISCUSSION. 

The  appearance  and  subsequent  behavior  of  bud  variations  in  Coleus 
present  numerous  analogies  to  various  phenomena  of  variation  exhib- 
ited by  members  of  a  seed  progeny  of  hybrid  origin. 

In  respect  to  the  definiteness  of  the  characters  contrasted,  green  and 
yellow,  red  and  non-red,  the  color  patterns  arising  by  bud  variations 
are  as  different,  at  least  in  their  extremes,  as  one  could  expect  in  the 
members  of  a  seed  progeny  even  of  hybrid  origin.  This  is  especially 
noticeable  in  comparison  with  the  seed  progenies  of  Coleus  itself.  The 
solid-red  epidermis  and  the  no-red  epidermis  represent  two  extremes 
fully  commensurate  with  the  so-called  presence  and  absence  of  a 
character  and  the  bud  variations  giving  these  were  fully  as  different  as 
the  types  arising  in  seed  progeny.  The  intermediates  red  blotched  and 
solid  red  upper  center  are  pattern  characters  equally  distinct  both  in 
manner  of  appearance  and  in  vegetative  constancy.  The  same  is  true 
of  the  extremes  of  development  of  both  green  and  yellow. 

Plants  of  the  same  pattern  in  a  single  line  of  descent,  both  in  the 
same  generation  and  in  successive  generations,  frequently  produced 
the  same  type  of  variation  independently,  a  behavior  quite  analogous 
to  the  segregations  that  reappear  in  each  generation  of  a  hybrid  line 
or  in  the  successive  generations  of  the  progeny  of  a  mutating  plant. 
Many  of  these  variations  show  a  return  to  a  parental  pattern,  just  as 
a  recessive  parental  quality  reappears  as  a  result  of  segregation  in 
hybrid  progeny.  There  is  much  in  such  reappearance  of  patterns  that 
is  quite  identical  with  the  phenomena  of  alternative  inheritance. 

Two  plants  identical  in  appearance  and  derived  from  adjacent 
branches  on  the  same  plant  may  give  quite  different  progenies  in  suc- 
sessive  generations.  One  line  may  be  very  constant  and  uniform,  the 
other  may  give  numerous  bud  variations  of  wide  range.  This  is  a 
familiar  phenomenon  in  hybrid  seed  progenies  where  certain  of  the 
plants  of  any  generation,  although  apparently  identical,  give  quite 
different  progenies.  In  Mendel's  experiments  with  Pisumjor  example, 
although  the  F2  yellow  peas  of  the  cross  green  X  yellow  were  similar, 
they  gave  different  progenies.  Some  produced  only  yellow  peas,  while 
the  progeny  of  others  gave  both  yellow  and  green. 

In  the  strains  of  Coleus  studied  b}^  the  writer,  certain  types  of 
bud  variation  occur  more  frequently  than  others.  Loss  of  yellow  was 
more  frequent  than  loss  of  green,  and  loss  or  decrease  of  red  was 
more  frequent  than  increase  of  red.  In  the  entire  series  of  plants 
derived  by  vegetative  propagation  there  was  decided  predominance  of 
green  over  yellow,  of  red  blotched  and  no  red  over  solid  red,  of  the  blumei 
character  of  leaf-base,  and  of  the  shallow  or  crenate  lobing  of  the 
leaves.  These  same  characters  show  marked  tendencies  for  domi- 
nace  among  the  members  of  the  seed  progeny. 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  59 

Such  phenomena  of  variation  appearing  in  hybridization  experiments 
are  usually  considered  as  due  to  segregation  and  recombination  of 
hereditary  units  during  the  processes  of  self-  or  cross-fertilization. 
Bud  variations  in  vegetatively  propagated  plants  are,  of  course,  inde- 
pendent of  such  recombinations. 

That  bud  variations  are  generally  due  to  a  complete  loss  during  cell 
division  is  not  substantiated  by  the  results  here  reported  or  by  the 
bulk  of  other  experimental  work.  In  the  majority  of  cases  the 
character  concerned  does  not  breed  true.  Mendelian  students  have 
interpreted  this  to  mean  that  such  bud  variations  are  produced  by  a 
loss  of  only  one  factor  of  a  diploid  pair,  giving  heterozygocity.  This 
illustrates  the  tendency  of  the  Mendelian  interpretation  to  assign  the 
numerous  cases  of  fluctuation  in  characters  to  heterozygocity  rather 
than  to  fluctuations  in  a  factor  or  to  irregular  mutational  changes 
spontaneous  in  the  organism. 

In  regard  to  the  range  of  expression  in  a  single  plant  the  laciniate 
leaf-shape  is  a  more  striking  character  even  than  the  color  patterns. 
It  arose,  as  already  noted,  in  13  individuals  obtained  by  vegetative 
propagation,  but  these  were  all  derived  from  a  few  plants  of  the  next 
preceding  generation.  This  character  was  inherited  through  vegeta- 
tive propagation  by  all  plants  grown  but  one,  but  the  leaves  on  each 
individual  plant  varied  from  deeply  laciniate  to  fully  entire.  Plants 
raised  from  seed  gave  all  types  from  extremely  laciniate  to  fully  entire, 
the  particular  type  appearing  in  the  first  leaves  that  developed  and 
remaining  quite  constant  for  all  leaves  developed  in  the  6  months  that 
the  plants  have  been  grown.  The  special  point  of  interest  is  that  a 
single  individual  of  the  laciniate  group  passed  through  a  series  of 
fluctuations,  giving  all  grades  of  leaf-shape  from  entire  to  fully  laciniate. 
The  range  in  a  single  plant  is  greatly  more  marked  than  the  dilTerences 
between  the  Urtica  hybrids  (Correns,  1905, 1912),  in  which  the  serrated 
type  of  leaf  was  dominant.  In  the  hybrids  of  the  normal  and  the 
laciniate  types  of  Chelidonium  majus  (de  Vries,  1900)  there  seems  to  be 
no  published  data  regarding  the  range  of  variation  in  the  F2  generation. 
Of  the  hybrids  between  palm-leaf  and  fern-leaf  types  of  Primula  sinen- 
sis, Bateson  (1909)  states  that  ''dominance  is  usually  complete," 
but  that  he  has  seen  two  strains  with  intermediate  leaf-shape.  Greg- 
ory (1911)  states  that  'Hhe  palmate  character  is  dominant,  though  a 
slight  difference  can  sometimes  be  recognized  between  pure  and  hetero- 
zygous palmate  types."  Crosses  between  an  ivy-leaf  (a  palmate 
shape  with  margins  crenate)  with  the  fern-leaf  gave  the  normal  palmate 
leaf  as  an  Fi  hybrid.  The  F2  generation  exhibited  a  wide  variation, 
which  Gregory  groups  into  4  classes  and  assumes  that  shape  of  the 
entire  leaf  and  crenation  of  the  margin  are  two  independent  characters. 

On  the  basis  of  character  of  lobing  of  climax  leaves,  Shull  (1911) 
distinguishes  four  biotypes  in  Bursa  hursa-pastoris.     The  view  that 


60  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

hybrid  forms  always  segregate  out  into  only  these  types  is  somewhat 
in  doubt,  for  Shull  (p.  9)  finds  that  a  plant  classed  as  simplex  gave 
unexpectedly  a  mixed  progeny  with  defective  ratio,  so  that  it  is  clear 
that  the  assumed  hereditary  "gene"  became  less  potent.  Hus  (1914) 
distinguished  in  a  culture  of  Capsella  hursa-pastoris  4  forms  different 
from  those  of  Shull  and  added  another  factor  which  he  considers 
determines  the  narrow  character  of  early  leaves  in  certain  forms. 

In  none  of  these  studies  have  the  individuals  of  the  F2  exhibited 
greater  variation  among  themselves  than  have  single  plants  of  Coleus 
with  the  laciniate  leaf,  nor  have  the  individual  parent  plants  been 
more  distinct  and  uniform  as  a  plant  than  the  plants  derived  from 
seed  progeny  of  Coleus,  and  grown  for  a  period  of  several  months. 
Furthermore,  there  has  been  evidently  no  attempt  to  select  persistently 
intermediate  types  for  modified  potency  of  characters.  Furthermore, 
emphasis  has  not  been  laid  to  selection  of  variations  in  a  seed  progeny 
of  a  single  individual  or  in  a  line  propagated  vegetatively.  The 
more  intensive  Mendelian  studies,  such  as  those  by  Shull,  Hus,  and 
Gregory,  indicate  that  the  character  of  leaf-shape  is  complex  and  that 
selection  studies  along  the  lines  indicated  ma\'  reveal  further  data 
on  variations  in  the  potency  of  characters. 

The  historical  evidence  and  the  studies  of  seed  progeny  reported 
above  indicate  that  the  strain  of  Coleus  studied  is  most  certainly  of 
mixed  parentage.  If  this  strain  had  been  studied  solely  in  its  seed 
progeny  the  variations  obtained  would  be  attributed  by  many  modern 
geneticists,  I  venture  to  imagine,  to  chance  combinations  of  hereditary 
units.  Yet,  as  has  been  fully  reported  above,  these  variations  are 
analogous  to  and  even  identical  in  nearly  all  cases  with  those  arising 
by  bud  variation. 

At  this  point  we  may  note  that  modern  genetics  have  furnished  no 
evidence  as  to  the  real  nature  of  the  characters  considered.  What 
we  may  call  the  genetical  or  breeding  value  of  characters  has  alone 
been  emphasized.  Characters  have  been  considered  solely  in  regard 
to  their  unity  of  expression  in  successive  generations  of  plants  of 
selfed  or  hybrid  origin.  Special  emphasis  has  been  placed  on  the 
reappearance  of  characters  and  upon  their  phylogenetic  significance. 
From  this  standpoint  we  may  further  consider  specifically  the  char- 
acters concerned  in  these  studies  of  Coleus. 

Considering  first  the  color  characters,  we  note  that  the  pattern  of 
the  parent  plants  was  a  mosaic  of  green,  yellow,  red  (or  blue),  and  non- 
red  cells.  In  the  mature  leaf  the  cells  are  apparently  qualitatively 
different,  and  furthermore,  the  color  differences  between  the  various 
cells  are  identical  with  the  color  differences  between  entire  leaves 
and  branches  derived  by  bud  variation.  The  contrast  between  pure 
j^ellow  and  pure  green  leaves  in  the  bud  sports  and  on  plants  of  seed 
origin  is  the  same  that  exists  between  green  and  yellow  cells  that  may 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  61 

be  adjacent  in  the  same  leaf.  Likewise,  the  differences  between 
solid-red  epidermis  and  the  non-red  epidermis  is  the  same  that  exists 
between  red  and  non-red  cells  in  the  epidermis  of  a  single  leaf  having  a 
red-blotched  pattern.  In  the  blotched  patterns  the  cells  of  the  epi- 
dermis are  either  red  or  non-red.  The  number  of  similar  cells  that 
are  adjacent  to  each  other  determine  the  size  of  the  blotches. 

The  facts  regarding  development  and  distribution  of  the  colors  raise 
sharply  the  question  whether  the  possibility  for  development  of  red, 
yellow,  and  green  is  possessed  by  every  cell  that  is  formed  or  whether 
these  possibilities  have  been  sorted  out  by  qualitative  divisions  in  a 
Weismannian  sense.  A  sharp  distinction  should  be  made  between  a 
character  that  is  metidentical  and  one  that  can  only  belong  to  a  group 
of  cells.  In  Detto's  (1907)  sense,  power  to  produce  green,  yellow,  and 
red  seems  to  be  metidentical;  that  is,  this  abiUty  may  be  strictly  a 
property  of  all  cells,  while  the  different  patterns  appearing  so  strikingly 
as  characters  of  entire  leaves  can  not  in  any  sense  be  metidentical. 

Most  important  evidence  regarding  the  development  of  leaves  with 
red-blotched  patterns  was  obtained  by  the  cytological  studies  conducted 
by  Mr.  E.  G.  Arzberger.  In  the  early  stages  of  leaves  that  later  will 
become  red-blotched,  all  the  epidermal  cells  possess  red  pigmentation, 
forming  a  complete  periclinal  layer  of  red  cells,  and  in  this  respect  the 
early  stages  of  leaves  with  solid-red  and  red-blotched  patterns  are  alike. 
In  solid-red  patterns  all  cells  continue  to  possess  red  sap  color,  while 
in  blotched  types  the  red  disappears  in  certain  cells.  The  evidence  is 
clear  in  these  cases  that  all  the  epidermal  cells  arise  from  cells  having 
red  cell  sap  and  in  this  respect  are  potentially  alike. 

The  differences  in  patterns  in  respect  to  red  are  largely  those  of 
quantity;  the  total  of  red  cell  sap  in  a  leaf  is  comparatively  large 
in  solid-red  patterns  and  much  less  in  blotched  patterns,  of  which  there 
is  every  grade  to  the  no-red  epidermal  patterns.  Differences  in  dis- 
tribution are  also  involved.  There  is  the  tendency  for  red  to  be  in  the 
epidermis  rather  than  in  the  subepidermal  tissues.  Usually  it  is  in 
both  upper  and  lower  surface,  but  in  one  pattern  (fig.  10)  the  red  is 
almost  entirely  massed  in  the  upper  epidermis  and  in  the  center  of  the 
leaf.  In  the  blotched  types  the  number,  size,  and  degree  of  coalescing 
of  the  blotches  present  every  gradation  from  a  finely  blotched  condition 
to  a  sohd  red. 

While  the  total  amoimt  of  red  and  its  distribution  in  the  epidermis 
determines  quite  definite  and  constant  patterns,  it  should  be  said  that  no 
plant  has  been  obtained,  either  in  seed  or  vegetative  propagation,  that 
was  entirely  devoid  of  red.  Subepidermal  cells  may  also  possess  red 
cell  sap;  red  sap  is  quite  pronounced  in  veins  of  leaves  (see  figs.  13 
and  14),  and  varying  amounts  can  be  seen  in  stems.  In  some  plants, 
especially  those  of  seed  origin,  no  trace  of  red  can  be  seen  except  at 
the  nodes,  and  here  the  amount  for  different  plants  varies  from  faint 


62  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

rings  to  quite  a  definite  band  or  zone  of  red  cells  extending  through  the 
stem.  Also  the  stems  of  plants,  especiallj^  those  with  blotched  patterns, 
may  have  large  irregular  streaks  of  red-colored  tissue  and  we  may  say 
are  internally  blotched.  In  general,  plants  with  a  decided  nodal 
zone  of  red  were  those  with  epidermis  free  of  coloration,  and  on  the 
other  hand,  plants  with  solid  epidermal  red  or  heavily  blotched  epider- 
mal red  had  as  a  rule  slight  localization  of  red  in  zones.  There  was  also 
the  marked  localization  of  red  in  upper  center  of  leaves  (see  fig.  10),  with 
almost  complete  absence  of  red  below,  although  in  many  leaves  of  this 
type  small  blotches  of  red  were  evident  on  the  under  side  (see  fig.  10a). 

All  these  conditions  indicate  that  the  possibility  of  producing  and 
possessing  red  cell  sap  is  a  specific  property  of  all  cells  and  that  the 
distribution  giving  localization  at  nodes,  in  streaked  areas  in  stems,  and 
in  subepidermal  tissues,  and  in  the  epidermis  either  as  a  uniform  red  or 
a  blotched  red  are  dependent  on  two  fundamental  conditions :  (a)  total 
amount  of  pigmentation,  and  (6)  the  appearance  of  it  in  certain  centers 
of  concentration.  The  facts  as  described  for  Coleus  seem  to  indicate 
the  such  conditions  are  determined  largely  by  intercellular  relations. 

Chemical  studies,  in  general,  show  that  differences  in  quality, 
quantity,  and  distribution  of  pigments  in  flowers  and  leaves  are  cor- 
related with  changes  in  quality,  amount,  or  distribution  of  any  one 
of  such  substances  as  chromogens  (glucosides, phenols,  tannic  acid,  etc.), 
oxydases,  enzymes,  oxygen,  etc. 

That  marked  changes  in  color  quite  comparable  to  those  I  have  de- 
scribed for  Coleus  may  result  from  slight  chemical  changes  is  well  shown 
by  the  chemical  studies  of  various  members  of  the  genus  Monarda. 
The  results  of  these  studies  are  fully  summarized  by  Wakeman  (1911). 
The  isolation  and  determination  of  yellow  and  red  pigments  and  a  study 
of  their  chemical  relatives  have  given  rise  to  the  quinhydrone  hypoth- 
esis of  plant  pigmentation.  It  is  considered  that  the  plant  oxidizes 
thymol  or  carvacrol  to  a  series  of  oxidation  products  of  yellow,  orange, 
and  red  colors,  but  all  closely  related  to  each  other.  Of  these,  hydro- 
thymoquinone,  thymoquinone,  and  dihydroxythymoquinone  have  been 
definitely  isolated  and  determined.  These  in  turn  have  the  capacity  of 
adding  phenols  yielding  highly  colored  phenoquinones  and  quinhy drones. 

Wakeman  (1911,  p.  Ill)  states: 

"Taking  into  consideration  only  those  compounds  that  have  been  isolated 
or  whose  presence  has  been  indicated  in  the  monardas  thus  far,  the  number  of 
possible  pigments  becomes  truly  bewildering." 

Furthermore,  some  of  the  pigments  are  phenolic  in  character  and 
can  combine  with  metallic  constituents  of  the  plants,  giving  rise  to 
different  shades  of  the  original  pigment. 

The  highly  colored  red  and  purple  pigments  of  the  stems  and  leaves, 
and  the  yellow  and  purphsh  pigments  of  the  flowers  in  Monarda  fistulosa 
are  thus  quite  definitely  identified  as  mixtures  of  quinhydrones  which 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  63 

are  shown  to  be  direct  products  of  the  plant.  The  important  point  is 
that  while  the  pigments  are  themselves  of  highly  complex  chemical 
substances,  the  changes  which  produce  marked  differences  in  color  are 
very  slight. 

It  is  also  significant  that  marked  differences  may  exist  between  the 
parts  of  a  single  flower.  The  simplest  of  color  patterns  in  Antirrhinum 
treated  by  Wheldale  (1914,  p.  110)  illustrates  this  condition.  Flowers 
of  the  yellow-flowered  variety  possess  a  pale  yellow  pigment  in  the  tube 
of  the  corolla,  a  deep  yellow  pigment  in  the  lips,  and  a  still  deeper 
patch  on  the  palate.  In  attempting  to  harmonize  these  varied  con- 
ditions with  a  Mendelian  factorial  analysis,  the  power  to  produce  these 
pigments  in  spatial  relation  is  assigned  to  a  factor  Y.  In  the  "ivory" 
flower  a  pale  pigment  is  found  in  the  tube  and  Ups  and  a  yellow  pigment 
only  in  the  epidermis  of  the  palate.  Power  to  produce  pale  pigment 
quite  generally  in  the  flower,  and  to  inhibit  the  formation  of  deep  yellow 
every^^here  but  in  the  epidermis  over  the  palate  is  assigned  to  a  single 
factor,  /.  The  conditions  of  color  distribution  are  in  themselves  color 
indicators  that  different  processes  occur  in  different  parts  of  the  same 
flower,  giving  different  kinds  of  substances  in  different  amounts  and 
with  different  distribution. 

Studies  of  oxydase  reactions  in  different  tissues  indicate  that  inter- 
cellular relations  are  of  much  importance  in  determining  the  distri- 
bution of  pigments.  By  means  of  micro-chemical  tests  Keeble  and 
Armstrong  (1912)  obtained  evidence  that  the  distribution  of  oxydase  in 
various  types  of  Primula  is  closel}^  correlated  with  the  development  of 
anthocyanin.  In  general  they  find  the  oxydase  most  abundant  in 
epidermal  and  in  vascular  tissue.  The  extent  of  ox3'dase  distribution 
differs  much  in  different  varieties.  They  make  the  highly  interesting 
observation  that  action  of  chloroplasts  seems  to  act  as  an  inhibitor 
of  oxidase  formation  or  of  the  production  of  chromogen.  Varied  types 
of  color  pattern  in  flowers,  and  even  anthoc^-anin  development  in 
stems  and  leaves,  is,  according  to  these  investigations,  closely  related 
to  distribution  of  oxydases  and  chromogens. 

It  is  of  further  interest  to  note  that  patterns  resulting  from  such 
qualitative  and  quantitative  reactions  which  depend  in  considerable 
degree  on  flow  of  substances  in  a  plant  may  be  quite  uniform  not  only 
in  the  flowers  of  a  plant,  but  among  the  various  members  of  its  seed 
progeny.  Such  phenomena  have  led  to  the  assumption  that  patterns 
are  represented  as  such  in  germ-cells  by  hereditar}-  units.  A^Tien,  how- 
ever, hybridization  occurs  between  varieties  having  even  the  simplest 
of  patterns,  the  F2  generation  more  often  than  otherwise  presents  a  most 
remarkable  range  of  types.  Usually,  this  sort  of  diversity  results  in 
crosses  betw^een  varieties,  especially  when  the  color  patterns  of  flowers 
and  leaves  are  concerned  and  gives  results  that  could  not  be  predicted 
with  any  degree  of  accuracy. 


64  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

Riddle  (1909),  in  a  comprehensive  survey  of  the  chemical  and  phys- 
iological facts  regarding  the  origin  and  nature  of  melanin  pigments, 
points  out  (p.  323)  that  "a  single  chromogen  acted  upon  by  a  single 
enzyme  usually  produces  several  colors  depending  upon  the  degree  of 
oxidation  involved."  He  gives  conclusive  evidence  that  "the  power  to 
oxidize  tyrosin  compounds  is  not  dependent  primarily  upon  germinal 
segregation,  but  rather  upon  active  tissues,  relations,  and  conditions," 
and  that  local  conditions,  especially  in  pathological  cases,  determine  the 
production  of  melanin.  In  general  Riddle  shows  the  inadequacy  of  a 
strict  interpretation  of  color  inheritance  in  animals  on  the  basis  of  unit 
factors  and  gametic  purity. 

In  the  light  of  all  the  chemical  studies  on  pigmentation,  it  seems 
clear  that  qualitative  reactions  are  concerned  which  involve  the  pro- 
duction, flow,  and  assembling  of  substances  through  the  relations  and 
interactions  between  cells.  That  these  interactions  should  be  so  widely 
readjusted  in  a  hybrid  progenj^  resulting  in  such  varied  expression  of 
color  in  quantity,  quality,  and  distribution  is  suggestive  that  funda- 
mental readjustments  may  occur  more  readily  with  characters  that  are 
dependent  on  cellular  interactions  than  with  those  that  are  strictly  meti- 
dentical.  As  already  pointed  out,  the  variations  in  Coleus  propagated 
vegetatively  give  numerous  patterns  differing  widely  in  regard  to  quan- 
tity and  distribution  of  the  pigments  concerned  and  present  the  same 
sort  of  phenomena  of  readjustment  seen  in  seed  progeny  of  hybrids. 

In  questions  relating  to  the  development  of  color  patterns,  the 
Liesegang  precipitation  phenomena,  especially  as  developed  by 
Gebhardt  (1912)  and  Kiister  (1913),  seem  to  me  most  illuminating.  By 
the  various  phenomena  associated  with  rhj^thinic  precipitation  and 
crystallization  of  mineral  solutions  in  gelatin  plates,  Gebhardt  was  able 
to  produce  simple  and  multiple  eye,  line,  and  flaked  patterns  strikingly 
similar  to  various  markings  in  butterfl}-  wings.  By  varjdng  the  sub- 
stances used,  modifjdng  the  amounts,  the  distribution,  the  degree  of 
concentration,  and  providing  for  interaction  between  areas  of  different 
concentration,  a  wide  range  of  markings  were  produced.  Gebhardt 
points  out  that  such  physical  and  chemical  phenomena  indicate  that 
the  distribution  of  pigment  even  in  the  intricate  markings  of  butterfly 
wings  may  be  due  to  an  epigenetic  regulation  of  the  quality  and  quan- 
tity of  such  substances  as  chromogen,  oxydase,  and  oxygen  and  the 
reciprocal  influences  of  different  centers  of  action.  It  is  pointed  out  that 
the  cell  boundaries  and  especially  the  position  of  veins  may  determine 
the  distribution  of  the  substances  involved  and  determine  the  relative 
locations  of  centers  of  action  as  well  as  areas  and  centers  of  no  action. 

Kiister  (1913)  extended  the  study  of  Liesegang  precipitation  pheno- 
mena to  the  effects  produced  in  capillary  tubes,  obtaining  various  types 
of  banded  precipitation  patterns  which,  as  he  emphasizes,  suggest  that 
similar  chemical  and  physical  processes  may  determine  many  tj-pes  of 
variegation  in  both  monocots  and  dicots. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  65 

In  Coleus  the  development  of  patterns  is  considered  by  the  writer 
to  be  due  largely  to  cellular  and  tissue  interactions  influencing  general 
and  metidentical  qualities  with  results  quite  analogous  to  the  Liesogang 
phenomena.  Changes  involving  red  are  on  this  basis  rather  simple 
cases  of  readjustment  influencing  the  total  amount  of  pigment  produced 
and  the  distribution  in  centers  or  areas.  The  ability  to  produce  the 
different  chemical  substances  concerned  with  the  final  development  of 
the  red  pigmentation  is  assumed  to  be  a  general  property  or  potentiality 
of  all  Coleus  cells.  The  assembling  of  all  the  products  necessary  for 
the  final  stages  in  its  development,  however,  are  determined  by  the 
amounts  produced  and  their  flow  to  centers  of  activity  and  interaction. 
The  development  of  red,  especially  in  the  subepidermal  tissues,  indi- 
cates that  this  is  the  case  and  suggests  strongly  that,  as  shown  by 
Overton  (1899),  changes  in  the  amounts  of  red  pigmentation  may  be 
closely  related  to  changes  in  the  sugar-content  of  the  sap.  In  Coleus, 
however,  it  is  clear  that  such  changes  arise  quite  spontaneously  in  the 
cells  and  tissues. 

In  respect  to  the  development  of  green  and  yellow  in  particular  cells, 
the  processes  seem  to  be  antagonistic.  Plastids  are  present  in  both 
green  and  yellow^  cells,  but  in  yellow  cells  they  are  fewer  in  number, 
smaller  in  size,  and  somewhat  distorted  in  shape.  The  green  and 
yellow  cells  are  subepidermal,  extending  from  upper  to  lower  epidermis. 
In  a  pure-green  leaf  all  these  cells  remain  green.  In  the  most  extreme 
cases  of  yellow  development  nearly  all  the  cells  fail  in  the  production  of 
chlorophyl.  The  different  patterns  result  from  variations  in  the 
relative  number  of  green  and  yellow  cells  and  in  the  grouping  of  the 
cells  of  like  color.  In  some  a  green  field  is  blotched  with  island-hke 
areas  of  yellow  cells,  in  others  the  central  area  of  green  is  bordered 
by  an  irregular  band  of  yellow,  and  again  the  yellow  may  be  situated  in 
the  center  with  a  green  band  at  the  border. 

In  a  leaf  with  blotched  or  banded  green-yellow  patterns  the  inter- 
mingled areas  of  green  and  yellow  cells  indicate  quite  clearly  that  both 
types  of  cells  are  derived  from  the  same  cells  in  the  growing-points. 
In  the  development  of  leaves  it  can  be  observed  that  while  the  yellow 
areas  are  in  evidence  when  the  young  leaves  unfold,  the  yellow  seems 
tinged  with  green,  and  that  as  the  leaf  grows  the  yellow  becomes 
more  intense.  Furthermore,  the  yellow  bleaches  until  in  old  leaves 
the  yellow  areas  change  to  a  pale  yellow  or  white,  while  the  green  areas 
are  still  bright.  As  a  leaf  dies  the  green  areas  become  pale  greenish 
yellow.  These  observations  indicate  that  many  of  the  cells  which 
later  become  yellow  are  actually  green  at  first. 

The  fluctuations  that  appear  substantiate  this  view.  A  plant  with 
yellow-bordered  leaves  ma}^  produce,  especially  in  winter,  new  leaves 
entirely  green,  and  thus  possess  for  some  time  old  leaves  of  green-yellow- 
red  blotched  pattern  and  younger  leaves  of  green-red  blotched  pattern. 


66  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

Later  the  leaves  formed  may  show  increased  amounts  of  yellow,  until 
by  midsummer  the  plant  is  uniform  for  the  green-yellow-red  blotched 
pattern.  Figures  14  and  14a  show  winter  and  summer  conditions  quite 
general  for  the  leaves  of  the  yellow-green  pattern,  which  indicates  that 
cells  may  be  either  green  or  yellow.  Furthermore,  all  degrees  of  varia- 
tion can  appear  as  bud  variations  affecting  segments  of  a  bud. 

Such  conditions  indicate  that  all  the  cells  are  potentially  green. 
If  this  be  accepted,  a  further  point  is  raised  regarding  the  source  of 
the  influences  leading  to  loss  of  green  and  to  development  of  yellow  in 
its  place.  In  respect  to  the  final  action  in  the  cells,  change  from  green 
to  yellow  is  itself  apparently  a  local  phenomenon  in  that  local  action  of 
plastids  is  affected. 

The  fluctuations  and  variations  in  the  extent  and  position  of  the 
green  and  yellow  tissues,  however,  indicate  that  here,  as  in  the  develop- 
ment of  red,  certain  intercellular  influences  are  operating.  The  con- 
figurations of  the  yellow  and  green  areas,  as  well  as  that  of  the  red,  are 
in  marked  degree  bounded  by  veins.  For  the  green  and  yellow  this 
is  most  strikingly  shown  in  figures  12  and  14.  For  the  red  similar 
conditions  are  seen  in  any  of  the  blotched  patterns,  as,  for  example, 
figures  2  and  5.  This  also  indicates  that  the  flow  of  substances  giving 
different  centers  of  distribution  and  concentration  is  the  important 
factor  in  the  production  of  patterns. 

In  respect  to  the  extent  and  degree  of  the  variations,  it  has  already 
been  pointed  out  that  no  plant  in  my  cultures  of  Coleus  has  been 
obtained  without  some  red  coloration  in  some  part  of  the  plant.  In 
regard  to  yellow,  however,  there  were  frequently  fluctuations  and  bud 
variations  giving  branches  with  no  yellow.  The  loss  of  yellow  appears 
to  be  complete  in  a  manner  that  suggests  loss  of  hereditary  qualities 
through  segregations,  but  even  in  constant  selection  of  pure-green  stock 
for  vegetative  propagation  about  half  of  the  offspring  show  return  to 
patterns  containing  yellow  both  by  fluctuations  and  by  bud  variations, 
with,  also,  cases  of  marked  spontaneous  appearance  of  yellow  in  a  few 
or  in  single  leaves.  To  say  that  the  power  to  produce  yellow  has  been 
latent  is  to  say  that  the  conditions  causing  its  development  can  arise 
in  an  apprently  spontaneous  manner. 

AVhile  the  evidence  indicates  that  the  ability  to  produce  green, 
yellow,  and  red  is  a  metidentical  property  of  the  cells,  it  is  equally 
clear  that  these  metidentical  properties  do  not  exist  as  units.  They  are 
subject  to  interaction  between  cells.  They  are  more  or  less  perma- 
nently modified  either  by  the  intercellular  relations  or  by  spontaneous 
intracellular  changes.  The  records  of  pedigree  show  very  clearly  that 
tendencies  to  give  vegetative  progeny  of  different  degrees  of  constancy 
and  variation  arise  or  exist  in  sister  branches  that  are  apparently 
quite  identical.  Such  tendencies  detected  by  the  pedigreed  cultures 
have  already  been  mentioned.     The  production  of  branches  which  give 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  07 

clones  of  plants  with  decreased  amounts  of  red  indicates  a  speoific 
decrease  in  power  to  develop  red  and  the  results  of  pedigree  culture 
indicate  that  continued  selection  in  this  direction  would  give  lines 
with  only  slight  amounts  of  red  or  perhaps  that  are  entirely  free  of 
all  red. 

The  metidentical  characters  of  green,  red,  or  yellow  are  themselves 
fluctuating  not  only  in  expression  but  in  inheritance  through  cell 
Hneage.  They  do  not  appear  to  be  independent.  I  have  never  yet 
obtained  a  plant  by  seed  or  by  vegetative  propagation  that  did  not 
possess  some  degree  of  red  coloration,  and  as  yet  no  strain  has  been 
isolated  that  was  pure  for  loss  of  yellow. 

On  the  whole,  vegetative  propagation  of  any  new  type  that  arises 
gives  a  progeny  that  exhibits  a  rather  marked  degree  of  constancy  with 
fluctuations  and  variations  about  a  new  mode.  Selections  for  pure 
green  do  not  give  a  progeny  of  pure-green  plants,  but  do  give  a  greater 
number  of  green  plants  than  does  any  other  pattern.  Selections  for 
decreased  red  or  for  increased  red  likewise  give  clones  with  this 
tendency  prevailing.  In  other  words,  readjustments  of  the  processes 
concerned  with  total  production  of  pigments  and  their  distribution 
tend  strongly  to  occur  in  growing-points  or  to  so  affect  them  as  to 
secure  a  certain  degree  of  permanency. 

The  assumptions  of  de  Vries  already  noted  in  the  introduction  seem 
to  apply  quite  adequately  to  the  behavior  of  the  metidentical  char- 
acters as  far  as  expression  in  individual  cells  is  concerned,  but  does 
not  fully  explain  the  phenomena  of  pattern  changes  as  well  as  the  con- 
ception of  a  further  influence  of  intercellular  relations,  modifications 
of  which  may  in  time  affect  more  or  less  permanently  the  expression  of 
metidentical  qualities.  x\ny  Mendelian  conception  of  pattern  factors 
that  are  units  in  heredity  is  quite  inadequate,  as  is  also  such  a  con- 
ception for  even  the  metidentical  characters. 

This  analysis  of  the  nature  of  variegation  and  the  significance  of 
bud  variation  in  Coleus  has  a  direct  bearing  on  the  nature  of  certain 
other  types  of  variegation. 

One  of  the  most  clearly  marked  types  of  variegation  is  that  of  the 
infectious  chlorosis.  The  best  known  cases  are  those  of  tobacco 
(Beijerinck,  1899;  Woods,  1899)  and  Abutilon  (Baur,  1904  and  1906; 
Lindemuth  1897,  1899,  1901,  1905,  and  1907).  All  investigations 
agree  that  in  these  types  the  variegation  is  not  transmitted  to  seed 
progeny.  The  searching  investigations  of  Beijerinck  and  Baur  lead 
to  the  conclusion  that  a  living  fluid  or  virus  carries  the  contagion.  In 
the  case  of  tobacco  the  infection  is  readily  accomplished  by  various 
agencies.  In  Abutiloti  grafting  is  necessary,  and  by  this  means  the 
variegation  has  been  transmitted  to  numerous  species  of  Abutilon  and 
related  genera.  Similar  types  of  infectious  variegation  exist  in  other 
groups  of  plants,  as  Fraxinus,  Jasminum,  Liburnum,  and  Ligustrum. 


68  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

In  the  infectious  variegation  of  the  type  seen  in  Abutilon  striatum 
thompsonii  the  pattern  is  a  mottled  one,  with  irregular  yellowish  areas 
mingled  with  the  green.  The  amount  of  yellow  varies  considerably, 
especially  according  to  intensity  of  illumination.  In  Abutilon  mega- 
potamicum  variegatum  especially  the  distribution  on  a  single  plant  is 
very  irregular  (see  Reid,  1914).  Only  a  few  blotches  may  appear  on 
a  leaf  and  often  entire  leaves  or  all  leaves  on  an  entire  branch  may 
be  pure  green.  Baur  (1906a)  found  green  branches  on  Abutilon  striatum 
thompsonii  and  was  able  fully  to  establish  that  they  were  immune. 

Immune  branches  arise  on  a  plant  as  bud  variations,  but  the  leaves 
differ  from  those  with  variegation  only  in  having  all  cells  immune,  for 
in  the  latter  a  part  of  the  cells  perhaps  remain  immune.  It  may  be 
noted  that  blotched  variegation  in  this  case  either  results  from  irregular 
immunity  or  to  irregular  distribution  of  virus,  and  hence  emphasizes 
the  intercellular  relations  concerned  with  distribution. 

V^Tiatever  the  nature  of  the  ''virus"  may  be,  it  is  fully  demonstrated 
by  Baur  that  it  is  a  product  of  the  diseased  cells  of  the  old  leaves  and 
is  transported  to  young  leaves  in  which  certain  areas  of  cells  succumb 
to  the  influence  while  others  do  not.  This  immunity  exhibited  by 
some  cells,  however,  may  suddenly  extend  to  entire  leaves  or  to  all  leaves 
on  a  branch.  As  to  the  flow  of  the  "virus,"  Baur  (1906a)  found  that 
it  could  pass  through  tissues  of  immune  strains  of  Abutilon  arboreum, 
causing  infection  to  non-immune  parts  beyond,  but  that  such  infection 
was  not  produced  if  immune  tissue  of  Lavatera  arborea  intervened. 

It  should  be  noted  that  this  type  of  variegation  can  not,  as  far  as  we 
know  now,  be  distinguished  by  appearance  from  other  types  of  mottled 
variegation.  Its  infectious  nature  and  the  failure  to  transmit  to  seed 
progeny  are  the  characteristics  of  these  cases.  As  noted  above, 
ShuU  (1914)  suggests  that  certain  yellow-flecked  types  of  variegation 
giving  very  irregular  transmission  to  seed  progenj^  may  be  of  such 
infectious  nature  that  it  can  be  carried  in  some  of  the  germ-cells. 
It  may  well  be  that  in  many  cases  of  variegation  (especially  of  the 
blotched  t>"pes),  the  disturbing  cause  producing  loss  of  chlorophyl 
may  be  quite  similar  in  nature  to  that  of  the  \agorously  infectious  types. 
In  the  latter  it  is  quite  clear  that  the  production  of  a  "virus"  in  varie- 
gated leaves  and  its  flow  to  3'oung  leaves  does  occur. 

Differences  in  the  extent  of  influence  of  such  a  "virus"  may  give  in 
some  types  an  apparent  inheritance  through  seed  progeny.  In  the 
infectious  types  we  know  nothing  definite  concerning  the  appearance 
of  variegation  in  the  plant  first  showing  it.  It  appears  (Reid,  1914) 
that  many,  if  not  all,  of  the  abutilons  with  infectious  variegation  arose 
through  grafting  with  one  original  strain,  Abutilon  striatum  thompsonii. 
The  presence  of  infectious  variegation  in  such  widely  differing  genera 
as  Nicotiana,  Fraxinus,  Abutilon,  and  Ligustrum  indicates  that  the 
condition  should  arise  spontaneously.     Frequent  and  almost  continued 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  69 

spontaneous  development  of  a  less  vigorous  virus  may  be  very  common, 
and  many  cases  of  variegation,  even  of  those  that  are  apparently  seed 
constant,  may  be  due  to  such  a  condition. 

Lindemuth  (1905)  determined  that  the  variegation  in  Coleus  was 
not  infectious  like  that  of  Ahutilon.  Just  what  types  he  used  is  not 
clear  from  the  data  given.  Conclusive  evidence  regarding  this  point 
has  not  been  obtained  by  the  writer.  Thus  far  grafting  experiments 
between  green-yellow  and  pure  green  types  have  shown  no  cases  of 
development  of  yellow,  and  the  writer  has  assumed  that  the  variegation 
is  not  at  least  vigorously  infectious. 

We  may  also  note  that  spontaneous  loss  of  ability  to  produce  green 
in  a  part  of  the  cells  of  the  growing-point  may  result  in  a  chimeral 
variegation  such  as  Baur  (1909)  reports  in  white-margined  types  of 
Pelargonium  zonale.  His  anatomical  studies  showed  that  in  growing- 
points  the  white-colored  tissues  may  lie  over  the  green,  forming  histo- 
genic  layers  and  giving  remarkable  permanency  of  the  pericUnal 
chimera  in  vegetative  propagation. 

In  maintaining  this  relation  white  cells  give  rise  to  white  cells  and 
green  to  green,  but  mechanical  readjustments  in  the  growing-points 
may  give  branches  with  quite  different  distribution  of  the  two  kinds 
of  cells.  Branches  may  thus  arise  with  sectorical  distribution  of  green 
and  white,  with  only  white  or  only  green  cells,  or  even  with  reversed 
positions  (Baur,  1909;  Stout,  1913).  These  readjustments  give  no 
new  qualities  to  cells,  nor  do  they  appear  to  involve  changes  of  either 
kind  to  the  character  of  the  other.  Yet  the  occurrence  of  numerous 
types  of  variegation  with  this  chimeral  relationship  indicates  that  such 
spontaneous  loss  is  not  infrequent. 

Certain  types  of  bud  variation  in  Coleus  present  features  quite 
similar  to  the  readjustments  that  appear  in  Pelargonium,  and  raise  the 
question  whether  there  is  possibility  of  spatial  readjustments  of  dis- 
tinct tissue  elements.  The  sudden  and  apparently  complete  loss  of 
epidermal  red  suggests  that  this  layer  may  exist  as  peripheral  in  a  more 
or  less  chimeral  relationship,  but  the  development  of  red  and  non-red  in 
the  adjacent  cells  of  the  epidermis  of  a  single  leaf  indicates  clearly  that 
these  differences  can  arise  within  cells  of  the  same  immediate  progeny. 
If  for  any  reason  a  part  of  the  epidermal  cells  fail  to  develop  red,  the 
red  might  be  absent  in  the  entire  epidermis  for  the  same  reason. 

The  loss  of  yellow  giving  pure  green  might  seem  to  be  due  to  the 
exclusion  of  yellow  cells.  Also,  the  sharp  contrast  between  patterns 
of  green-yellow  (figs.  2,  8,  and  12)  and  pattern  of  yellow-green  (figs.  6, 
11,  and  14)  suggest  the  possibility  of  a  spatial  readjustment  in  the 
growing-point  of  two  distinct  cell  elements.  But  certain  plants  in 
all  these  patterns  have  fluctuated,  getting  greener  to  a  green-yellow 
blotched  pattern  during  winter  and  returning  to  the  type  pattern  in  the 
next  summer,  and  often  exhibiting  at  one  time  in  a  single  row  of 


70  ESTABLISHMENT    OF   VARIETIES   IN    COLEUS 

leaves  various  gradations  between  the  extremes.  The  extreme  yellow 
types  show  some  green  areas  of  tissue.  The  extreme  green  type,  which 
appears  to  have  no  j^ellow,  gives  numerous  fluctuations,  and  cases  of 
well-marked  spontaneous  development  of  yellow.  The  distribution 
of  green  and  yellow  does  not  in  any  pattern  show  anatomically  a 
chimeral  distribution,  as  both  colors  are  much  intermingled  in  the 
subepidermal  tissue. 

That  a  certain  degree  of  chimeral  relationship  exists  in  certain 
patterns  of  Coleus  is  evident.  In  the  patterns  with  solid-red  epidermis 
the  epidermal  layer  is  rather  specialized  in  respect  to  concentration 
of  red  cell  sap.  These  apparent  chimeral  relationships  in  Coleus  are 
due  to  intercellular  development  of  patterns  rather  than  to  specific 
and  qualitative  differences  in  cells  as  such. 

Numerous  cases  of  variegation  are  induced  by  environmental  con- 
ditions. Cramer  (1907,  chap,  xi)  summarizes  cases  of  the  influence 
of  parasites,  of  soil  conditions,  light,  and  temperature  in  producing 
certain  types  of  chlorosis  and  variegation.  These  are,  we  may  say, 
direct  reactions  to  external  conditions,  which  in  most  cases  are  quite 
apparent.  At  first  thought  this  class  may  seem  quite  distinct  from 
what  are  considered  as  true  hereditary  types,  but  the  difference  is 
chiefly  one  of  degree,  for  there  are  few  types  of  cases  of  variegation 
that  do  not  fluctuate  in  response  to  certain  environmental  conditions. 
The  infectious  types  of  variegation  fluctuate  verj^  much  according  to 
degree  of  illumination  and  may  entirely  disappear  from  a  plant  if  it 
is  kept  in  darkness  for  sufficient  length  of  time.  The  types  with  green 
and  white  as  periclinal  chimeras  show,  perhaps,  least  fluctuation  in 
regard  to  environmental  influence. 

In  this  respect  what  I  have  called  fluctuations  in  Coleus  are  of  inter- 
est. Fluctuations  in  amount  of  red  present  in  blotched  types  is  con- 
stantly occurring.  One  plant,  at  the  present  writing,  has  some  branches 
with  leaves  sparsely  blotched,  as  in  figure  5,  others  with  the  red  blotches 
strongly  coalescing,  as  in  figure  29,  and  still  others  with  nearly  a  sohd 
epidermal  red.  These  fluctuations  do  not  seem  related  to  external 
environmental  conditions.  In  the  amounts  of  green  and  j-ellow,  the 
various  patterns  possessing  these  two  elements  showed  a  strong  ten- 
dency to  be  more  green  in  winter  and  more  yellow  in  summer.  The 
degree  of  fluctuation  was,  however,  not  uniform  for  the  different  sub- 
clones, for  the  several  plants  of  a  single  generation,  or  even  for  all  the 
branches  of  a  single  plant.  A  few  plants  maintained  a  maximum  of 
yellow  in  their  leaves  throughout  the  winter.  Still  it  is  very  clear  that 
many  cells  in  the  leaves  are  green  in  winter  which  would  have  been 
yellow  had  the  particular  leaves  developed  during  summer. 

In  certain  plants  variegation  appears  periodically.  These  types 
are,  perhaps,  in  a  strict  sense,  to  be  classed  with  the  preceding,  but 
differ  at  least  in  showing  marked  periodicity  which  may  well  be  due 


BY    THE    SELECTION    OF    SOMATKJ    VARIATIONS.  71 

in  large  measure  to  inner  conditions.  Cramer  (1907,  p.  128)  sum- 
marizes numerous  cases  of  those  showing  marked  differences  in  beha- 
vior. Certain  varieties  of  Quercus  have  pure-green  foliage  each  spring, 
but  later  produce  variegated  leaves.  Uhnus  scabra  var.  viminalis  is 
yellow  during  midsummer,  but  pure  green  at  close  of  the  summer. 
Linaria  biennis  is  pure  green  during  the  first  year  of  growth,  but  varie- 
gated in  the  second. 

These  phenomena  illustrate  again  that  cells  of  the  same  lineage  may 
fluctuate  in  development  from  a  maximum  of  green  to  various  degrees 
of  loss  of  chlorophyl,  with  often  development  of  yellow  coloration,  and 
that  such  phenomena  may  reappear  with  marked  constancy  in  progeny. 

Aside  from  these  classes  there  is  a  wide  range  of  types,  including 
many  cases  in  which  the  variegation  reappears  more  or  less  generally 
and  constantly  in  the  individual  plants  and  in  the  variety.  The 
variegation  appears  to  be  inherited,  at  least  in  a  certain  degree. 

In  the  propagation  of  variegated  plants,  much  general  data  has  been 
obtained  regarding  the  degree  of  constancy  both  in  vegetation  and  seed 
propagation.  Cramer  (1907,  p.  129)  gives  a  summary  of  numerous 
cases  where  the  variegation  disappears  in  certain  methods  of  vegetative 
propagation.  In  Cornus  mascula  variegata  (T.  M.,  Gard.  Chron., 
32:  952),  root-cuttings  give  pure-green  individuals,  but  plants  from 
layers  retain  the  variegation.  Such  cases,  however,  should  be  investi- 
gated wdth  special  regard  to  the  nature  of  the  variegation.  In  many  of 
the  cases  noted,  especially  those  in  Pelargonium,  the  variegation  may  be 
chimeral,  and  when  root-cuttings  are  made  the  green  cells  have  greater 
power  of  regeneration.  The  inconstancy  exhibited  in  vegetative  propa- 
gation, however,  is  no  greater  or  more  marked  than  that  which  develops 
on  a  single  individual.  Exact  evidence  of  the  inheritance  and  con- 
stancy of  variegation  through  pedigreed  vegetative  progenies  seems  to 
be  lacking.  \ATiile  in  some  varieties  the  variegation  appears  to  be 
quite  constant,  in  others  it  is  widel}^  variable. 

The  Mendelian  studies  that  have  been  made  of  the  seed  progeny  of 
variegated  plants  and  of  the  bud  variations  which  involve  changes  of 
pattern  show  likewise  a  wide  range  of  behavior.  Hybridization  studies 
involving  variegation,  as  has  been  pointed  out  in  the  introduction, 
indicate  clearly  the  wide  range  of  variability,  and  w^hat,  from  the 
Mendelian  viewpoint  of  unit  characters  or  unit  factors,  is  most  erratic 
inheritance.  These  studies  have  contributed  interesting  and  valuable 
data  on  the  sort  of  variations  one  may  espect  in  hybridization  studies 
of  this  kind,  but  they  indicate  verj^  clearly  that  the  assumed  factors  are 
themselves  fluctuating.  In  these  studies  the  variabilit}-  of  plants  used 
as  seed  parents  has  not  been  determined  by  vegetative  propagation. 
This  is  not  possible  in  all  cases,  but  whenever  it  is  possible  the  emphasis 
should  be  placed  on  this  line  of  investigation  if  one  is  to  speak  with 
certainty  regarding  the  nature  of  the  inheritance. 


72  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

If  we  turn  our  attention  to  the  various  phenomena  associated  with 
the  laciniate  character  of  leaves  and  petals,  we  find  the  same  degree  of 
irregularity  and  diversity  of  origin,  expression,  and  inheritance  that  is 
exhibited  by  variegations.  Cramer  (1907,  chap.  21)  devotes  a  chapter 
to  an  excellent  survey  of  the  facts  regarding  the  behavior  of  the  char- 
acter. It  is  interesting  to  note  that  Cramer  observes  that  varieties 
quite  constant  in  seed  progeny  are  likewise  very  constant  in  vegetative 
propagation.  There  are  numerous  cases  known  of  the  spontaneous 
development  of  a  laciniate  type  from  one  with  entire  leaves,  both  as 
seed  mutations  and  as  vegetative  mutations;  likewise  of  return  to  the 
types  with  entire  leaves.  While  some  cases  are  quite  constant,  others 
are  widely  fluctuating,  even  exhibiting  a  marked  periodicity. 

The  behavior  of  this  character  in  Coleus  is  most  striking  in  its  varia- 
bility of  expression  and  of  its  inheritance  as  a  periodic  variation  through 
vegetative  propagation. 

Fundamentally,  the  processes  involved  in  the  development  of  leaf- 
shape  are  quite  different  from  those  involved  in  the  production  of 
pigments  such  as  green,  yellow,  and  red.  The  shape  of  the  leaf  in 
general  depends  on  the  rate,  number,  and  regularity  of  cell  divisions 
in  the  different  planes  of  growth.  It  would  seem  that  a  general  and 
quite  uniform  series  of  cell-divisions  would  give  a  leaf  of  more  regular 
outline,  and  that  if  the  cell-divisions  in  the  growing  leaf  occur  irregu- 
larly, giving,  so  to  speak,  points  or  lines  of  more  rapid  growth  somewhat 
analogous  to  apical  growth,  with  a  more  or  less  multiple  dichotomy, 
then  cut,  lobed,  or  laciniate  leaves  would  result.  The  shape  of  the  leaf, 
it  would  seem,  is  determined  by  intercellular  relations  concerned  with 
the  manner  of  cell-divisions. 

A  comparison  of  the  variability  that  develops  in  vegetative  propa- 
gation with  that  occurring  in  the  seed  progeny  reveals  some  essential 
differences  between  the  inheritance  of  the  characters  involved.  While 
the  range  of  variation  is  quite  the  same,  there  is  a  marked  difference 
in  what  we  may  call  the  intensity  of  variation.  In  vegetative  propaga- 
tion the  degree  of  the  intensity  was  low,  with  reference  to  the  frequency 
of  the  appearance  of  new  color  patterns  or  to  the  development  of  the 
laciniate  character.  A  large  number  of  plants  were  grown  by  vege- 
tative propagation.  The  bud  variations  were  comparatively  infrequent, 
occurring  something  like  once  for  every  10  plants  grown,  giving  in  the 
course  of  6  generations  the  different  types  of  pattern  described.  In  a 
seed  progeny,  however,  practically  the  entire  range  of  variation  which 
appeared  in  vegetative  propagation  was  seen  in  a  single  seed  genera- 
tion comprising  50  plants.  The  processes  concerned  with  reduction  and 
fertilization  increased  the  intensity  of  variation  and  brought  out  in  a 
single  progeny  of  no  great  number  the  full  extent  of  variations. 

Color  patterns,  which  are  intercellular  characters  and  in  a  sense 
vegetative  types,  are  inherited  through  vegetative  propagation  in 
marked  degree,  while  in  seed  propagation  there  is  no  evidence  that  they 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  73 

are  inherited  as  such.  The  intercellular  relations  involving  amounts 
and  distribution  of  pigments  are  widely  and  suddenly  disorganized 
and  readjusted  during  the  processes  concerned  with  seed  formation. 
On  the  other  hand,  the  metidentical  characters  green,  yellow,  and  red 
are  quite  uniformly  transmitted  in  both  sorts  of  cell-divisions.' 

The  behavior  of  the  laciniate  character  is  significant  in  this  con- 
nection. After  this  character  made  its  appearance  in  a  plant  it  was  a 
constant  feature  in  the  development  of  all  plants  (but  one)  derived 
by  vegetative  propagation.  The  seed  progeny,  however,  exhibited 
wide  differences,  ranging  from  deeply  laciniate  through  all  degrees  or 
grades  to  the  entire  type  of  leaf.  Furthermore,  the  laciniate  leaf 
appeared  in  all  seed  progenies  thus  far  grown,  even  when  derived  from 
plants  of  a  Hne  in  which  the  character  had  never  appeared.  The  proc- 
esses of  reduction  and  self-fertihzation  in  a  Coleus  plant  seem  to  bring 
out  latent  possibilities  for  various  developments  of  leaf-shapes. 

It  remains  to  be  seen  if  any  of  the  types  appearing  in  seed  progeny 
are  more  constant  in  vegetative  or  in  seed  propagation  than  are  the 
similar  types  that  develop  by  bud  variation.  Already  several  bud 
variations  have  appeared  in  plants  of  seed  progeny,  indicating  vegeta- 
tive changes  in  the  processes  concerned  with  pigment  and  pattern 
formation.  The  colors  involved  in  the  variegation  of  Coleus  represent 
every  type  of  coloration  (green,  yellow,  white,  and  red  or  blue)  con- 
cerned with  the  variegation  and  coloration  of  plants.  There  is  no 
evidence  that  the  essential  nature  of  these  characters  in  Coleus  differs 
from  that  of  the  characters  concerned  with  variegation  and  pigmenta- 
tion in  corn  (Emerson,  1914) ,  in  Mirabilis  (Correns,  1909) ,  in  Melandrium 
(Shull,  1914), in  Pelargonium  (Baur,  1909),  in  Antirrhinum  (Baur,  1910). 

The  explanations  here  given  regarding  the  spontaneous  variability 
of  the  characters  concerned  in  the  development  of  pigments  and  of  the 
changes  in  intercellular  and  intertissue  relations  influencing  develop- 
ment of  color  patterns  in  Coleus  apply  equally  well  to  such  cases  as 
those  just  noted.  The  evidence  indicates  that  the  characters  in  ques- 
tion, and  most  especially  the  pattern  characters,  are  not  represented  by 
units  or  factors,  unless  these  assumed  factors  are  to  be  considered  in  a 
general  sense  as  temporary  conditions  descriptive  of  tj'pes  of  develop- 
ment and  not  as  particular  localized  units  of  germ-plasm,  which  is  the 
conception  that  gave  the  Mendelian  interpretation  its  definiteness  and 
simplicity. 

The  knowledge  of  the  nature  and  the  heredity  of  color  character- 
istics will  be  advanced  more  by  studies  of  the  natural  variability  of 
the  characters  involved  and  by  chemical  and  phj'sical  investigations  of 
the  processes  concerned  in  the  formation  and  distribution  of  such  sub- 
stances as  melanin,  flavone,  and  phenol  compounds  rather  than  by 
further  elaboration  of  complicated  formulae  involving  multiple  factors 
that  attempt  to  explain  fluctuations,  inherited  variations,  and  cases 
of  increased  variability  that  appear  in  hybrid  seed  progenies. 


74  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

SUMMARY. 

(1)  A  single  variety  of  Coleus  propagated  vegetatively  by  cuttings 
in  two  main  clones  has  shown  (a)  gradual  fluctuations  and  (6)  sudden 
mutations,  giving  a  total  of  16  distinct  and  characteristically  different 
color  patterns. 

(2)  (a)  A  total  of  15  patterns  (see  diagram  2)  arose  by  sudden  muta- 
tion affecting  a  part  of  a  leaf  or  a  branch,  or  a  series  of  associated  leaves 
or  branches;  (b)  6  of  these  15  patterns  (figs.  2,  4,  5,  10,  13,  and  13a) 
also  appeared  among  the  fluctuating  variations;  (c)  one  type  of  color 
pattern  (fig.  15)  has  thus  far  appeared  only  as  a  fluctuating  variation. 

(3)  (a)  Six  (diagram  2,  and  figs.  1,  4,  5,  6,  8,  and  12)  of  the  15  color 
patterns  arose  directly  from  the  parent  type  bj^  sudden  bud  variation. 
One  of  these  6  (fig.  4)  also  appeared  as  a  fluctuating  variation.  (6)  Five 
of  these  6  tj^pes  (figs.  4,  5,  6,  8,  and  12)  propagated  by  cuttings  showed 
further  fluctuations  and  bud  variations,  giving  (a)  the  parent  type 
(fig.  2),  (6)  4  of  the  6  types  already  directly  derived  (figs.  1,  2,  5,  and  12), 
and  (c)  8  new  types  (figs.  9,  10,  11,  13,  14,  and  16,  also  8a,  13a,  not 
illustrated). 

(4)  The  variations  in  the  development  of  color  patterns  mentioned 
above  involve  (a)  increase  and  decrease  of  green  and  yellow,  (b)  increase 
and  decrease  of  red  pigmentation,  (c)  reversals  of  the  relative  positions 
of  the  green  and  yellow  by  which  a  type  with  green  center  and  yellow 
border  (fig.  2)  gave  one  with  yellow  center  and  green  border  (fig,  6), 
and  (d)  changes  in  the  distribution  of  the  red  pigmentation  especially 
giving  concentration  in  the  epidermis  of  the  upper  surface  of  the  leaves. 

(5)  Progeny  of  11  types  of  color  pattern  have  been  grown  through 
from  2  to  6  generations,  as  follows: 

Type  of  figure 1         2        4      5       6       8     9     10     12     13     14 

Number  of  generations 2         6        6      6       6       5     2      2       2      2      2 

Total  number  of  plants 11    337     198    90     41     54     8      7       7       7      4 

Some  of  these  types  have  shown  themselves  more  constant  than  the 
parent  type  (fig.  2),  others  were  less  constant.  All  varied  about  a  new 
mode  and  all  would  be  considered  good  horticultural  races. 

(6)  The  relative  constancy  of  color-pattern  types  derived  by  the 
accumulation  of  fluctuating  variations  was  tested  in  two  cases:  Type 
of  figure  2  (see  clone  14  of  table  2),  3  generations,  total,  45  plants; 
type  of  figure  4  (see  clone  13  of  table  3)  3  generations,  total,  79  plants. 
In  both  cases  the  constancy  of  the  progeny  showed  no  essential  differ- 
ence from  that  of  the  same  types  obtained  by  sudden  bud  variations. 

(7)  Variations  in  leaf  form,  even  more  striking  than  changes  in  color 
patterns,  from  entire  to  deeply  laciniate-leaved  forms,  appeared  in  13 
instances  as  fluctuations  affecting  an  entire  plant  and  in  one  case 
(during  the  winter  of   1914-14)   as  a  bud  variation.     The  striking 


BY    THE    SELECTION    OF    SOMATIC    VARIATIONS.  75 

feature  of  this  variation  is  the  marked  tendency  to  the  production  of 
laciniate  leaves  during  winter  or  after  a  period  of  particular  drought 
during  summer. 

(8)  The  behavior  of  the  laciniate  character  was  observed  in  three 
subclones  (see  table  4)  of  4,  2,  and  2  generations,  totaling  68  plants. 
In  67  of  these  plants  the  laciniate  type  of  leaf  appeared  as  a  definite 
character,  with  a  marked  tendency  to  a  periodic  summer  and  winter 
fluctuation, 

(9)  Plants  with  the  laciniate  leaf  character  also  showed  wide  fluctu- 
ations in  regard  to  the  degree  of  green  or  yellow  coloration.  When 
grown  for  a  period  of  a  year  from  cuttings  made  in  autumn,  the  leaves 
were  as  a  rule  entire  and  slightly  yellov/  in  autumn,  deeply  laciniate 
and  pure  green  in  winter,  and  entire  and  very  yellow  during  the 
following  summer. 

(10)  In  sexual  reproduction  all  the  principal  types  of  variegation  and 
leaf-shape  appear  at  once  in  an  Fi  generation,  with  also  numerous 
types  that  were  intermediate  and  fluctuating.  The  extremes  of  vari- 
ation are  no  greater  than  those  obtained  in  vegetative  propagation, 
although  some  new  types  of  entire  leaves  were  thus  obtained. 

(11)  Between  any  two  types  numerous  intermediates  arose,  showing 
that  we  have  here  no  evidence  of  the  somatic  segregation  of  invariable 
pattern  factors. 

(12)  In  the  bud  variations,  decrease  of  red  occurred  with  about  twice 
the  frequency  as  did  increase  of  red;  likewise  decrease  of  yellow 
occurred  about  twice  as  often  as  the  increase  of  yellow,  indicating  a 
definite  tendency  for  variations  in  the  direction  of  the  increase  of  green 
and  the  decrease  of  red.  These  facts  are  doubtless  due  to  fundamental 
relations  between  the  chemical  compounds  involved. 

(13)  The  types  of  color  changes  involving  (a)  green  and  yellow  and 
(6)  red  and  non-red  occurred  entirely  independently  of  each  other. 

(14)  The  types  produced  by  bud  variations  are  the  equivalents  of 
the  "Kleinarten"  or  the  ''biotypes"  commonly  occurring  in  cultivated 
species  propagated  by  seed. 

(15)  Selection  within  clones  is  effective  in  securing  progenies  of  new 
types  with  as  high  degrees  of  constancy  as  is  possessed  by  ordinary 
cultivated  races. 

(16)  The  results  indicate  that  slight  variations  arising  either  as 
sudden  mutations  or  as  gradual  fluctuations  can  perpetuate  themselves. 

(17)  The  green,  yellow,  red,  and  non-red  colorations  in  Coleus  can 
best  be  characterized  as  metidentical  characters;  that  is,  they  are  the 
same  in  the  cells  as  in  the  tissue  and  their  appearance  is  possible  in 
the  development  of  any  cell. 

(18)  The  distribution  of  the  colors  giving  pattern  characters  are 
properties  of  groups  of  cells  and  tissues  as  such.  Pattern  characters 
are  probably  due  entirely  to  tissue  and  cellular  interactions. 


76  ESTABLISHMENT    OF    VARIETIES    IN    COLEUS 

(19)  The  explanation  suggested  by  the  production  of  patterns  in 
colloids  by  the  Liesegang  precipitation  phenomena,  especially  as  applied 
by  Gebhardt  to  the  markings  of  butterfly  wings  and  by  Kiister  to  the 
development  of  many  types  of  variegation  in  plants,  seems  to  apply 
to  the  production  of  color  patterns  in  Coleus.  On  this  view  color  changes 
may  be  considered  as  due  to  the  formation  of  different  diffusion  centers 
for  the  development  and  concentration  of  pigments. 

(20)  Bud  variations  in  Coleus  are  (a)  common;  (6)  give  numerous 
different  types  which  may  be  vegetatively  quite  constant  from  the  first 
or  can  be  made  so  by  selection;  (c)  show  development  of  certain 
types  more  commonly  than  others;  (d)  produce  reversions  to  parental 
t3qDes;  (e)  give  development  of  different  degrees  of  variability  among 
sister  clones;  (/)  exhibit  spontaneous  changes  in  the  fundamental  color 
characters  (metidentical)  and  in  the  cellular  and  tissue  processes  result- 
ing in  color  patterns. 

(21)  The  results  show  that  in  Coleus  asexual  and  sexual  reproduction 
are  not  fundamentally  different  in  respect  to  the  extent  and  range  of 
variation. 

A.  B.  Stout, 

Director  of  the  Laboratories. 

New  York  Botanical  Garden, 

New  York  City,  February  10,  1915. 


BY   THE    SELECTION    OF    SOMATIC   VARIATIONS.  77 

BIBLIOGRAPHY. 

Andr6,  Ed.     1879.     Les  nouveaux  Coleus.     111.  Hort.,  26:  160. 

.     1880.     Coleus  blumei,  nov.  var.  hortenses.     111.  Hort.,  27:  52-53;  pi.  377. 

Bateson,  William.     1902.     Mendel's  principles  of  heredity. 

.     1909.     Mendel's  principles  of  heredity. 

.     1914.     Address  of  the  President  of  the  British  Association  for  the  Advancement 

of  Science.     Science,  ii,  40:  319-333. 
Bateson,  William,  and  R.  C.  Punnett.     1905.     A  suggestion  as  to  the  nature  of  the 

"wabiut"  comb  in  fowls.     Proc.  Cambridge  Phil.  Soc,  13:  165-168. 
Baur,  Erwin.     1904.     Zur  Aetiologie  der  infektiosen  Panachierung.     Ber.  Deutsch.  Bot. 

GeseU.,  22:  453-460. 
.     1906a.     Weitere  Mitteilungen  liber  die  infektiose  Chlorose  der  Malvaceen  und 

iiber  einige  analoge  Erscheinungen  bei  Ligustrum  und  Laburnum.     Ber. 

Deutsch.  Bot.  Gesell.,  24:  416-428. 
.     19066.     Ueber  die  infektiose  Chlorose  der  Malvaceen.     Sitz.-ber.  Preuss.  Akad. 

Wiss.,  1906:  11-29. 
.     1909.     Das  Wesen  und  die  Erblichkeitsverhaltnisse  der  "Varietates  albomargin- 

atae  Hort."  von  Pelargonium  zonale.     Zeit.  Ind.  Abs.  Vererbs.,  1:  330-.351. 
.     1910.     Untersuchungen  tiber  die  Vererbung  von  Chromatophorenmerkmalen  bei 

Malandrium,  Antirrhinum,  und  Aquilegia.     Zeit.  Ind.  Abs.  Vererbs.,  4: 

81-102. 
Beijerinck,  M.  W.     1899.     De  I'existence  d'un  principe  contagieux  vivant  fluide,  agent 

de  la  nielle  des  feuilles  de  tabac.     Arch.  N^erl.,  Ser.  ii,  3:  164-186. 
Blitme,  C.  L.     1826.     Plectranthus  scutellarioides  Bl.     Bijdragen  tot  de  Flora  van  Neder- 

landsch  Indie,  p.  837. 
Castle,  W.  E.     1912.     The  inconstancy  of  imit  characters.    Am.  Nat.,  46:  352-362. 
CoRRENS,  C.     1905.     Ueber  Vererbungsgesetze. 

.     1912.     Die  neuen  Vererbungsgesetze. 

.     1909a.     Vererbungsversuche    mit    blass    (gelb)    griinen    und    bunt    blattrigen 

Sippen  bei  Mirabilis  jalapa,  Urtica  pilulifera,  und  Lunaria  annua.     Zeit. 

Ind.  Abs.  Vererbs.,  1:  291-329. 
.     19096.     Zur  Kenntnis  der  RoUe  von  Kern  und  Plasma  bei  der  Vererbung.     Zeit. 

Ind.  Abs.  Vererbs.,  2:  331-340. 
Cramer,  P.  J.  S.     1907.    Kritische  Uebersucht  der  bekannten  Falle  von  Knospenvariation. 
Darwin,  Charles.     1868.     Variation  of  animals  and  plants  under  domestication.    2  vols. 
Detto,  Carl.     1907.     Die  Erklarbarkeit  der  Ontogenese  durch  materielle  Anlagen.     Biol. 

Centralb.,  27:  80-95,  106-112,  142-160,  162-173. 
de  Vries,  Hugo.     1889.     Intracellulare  Panagenesis. 

.     1900.     Das  Spaltungsgesetz  der  Bastarde.     Ber.  Deutsch.  Bot.  Gesell.,  18:  83-90. 

.     1901.     Die  Mutationstheorie. 

.     1913.     Gruppenweise  Artbildung. 

DoMBRAiN,  H.  H.     1867a.     Coleus  gibsonii.     Floral  Mag.,  6:  pi.  338  (with  description). 

.     1867b.     Coleus  veitchii.     Floral  Mag.,  6:  pi.  345  (with  description). 

East,  E.  M.     1908.     Suggestions  concerning  certain  bud  variations.     Plant  World,  11:  77. 
.     1910a.     The  transmission  of  variations  in  the  potato  in  asexual  reproduction. 

Contrib.  Lab.  Genetics,  Bussey  Inst.  Harvard  Univ.,  No.  III.     Biennial 

Report  Conn.  Exp.  Sta.,  1909-10,  119-161. 
.     19106.     A  Mendehan  interpretation  of  variation  that  is  apparently  continuous. 

Am.  Nat.,  44:  65-82. 
Emerson,  R.  A.     1914.     The  inheritance  of  a  recurring  somatic  variation  in  variegated 

ears  of  maize.     Research  Bull.  Exj).  Sta.  Neb.,  No.  4. 
Flammarion,  Camille.    1898.     Physical  and  meteorological  researches,  principally  on  solar 

rays,  made  at  the  Station  of  Agricultural  CUmatologj'  at  the  Observatory 

of  Juvisy.     U  S   Dept.  Agr.,  E.xp.  Sta.  Record.,  10:  103-114,  pi.  1. 
Gebhardt,  F.  a.  M.  W.     1912.     Die  Hauptzuge  der  Pigment  Verteilung  ira  Schmetter- 

lingsfliugel  in  Lichte  der  hiesegangschen  Niederschliige  in  Kolloiden. 

Verh.  Deutsch.  Zool.  Gesell.,  22:  179-204. 
Goodspeed,  T.  H.     1912.     Quantitative  studies  of  inheritance  in  Nicotiana  hybrids.     Univ. 

of  CaUf.  Publ.,  Bot.,  5:  87-168. 
Gregory,  R.  P,     1911.     Experiments  with  Primula  sinensis.    Jour.  Genetics,  1:  73-132; 

pi.  30-32. 


78  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS 

Herincq,  F.     1865.     Choix  de  plantes  a  feuillage  rouge.     Hort.  Frangais,  7:  308-311. 
Also  same  title  and  article  in  La  Belg.  Hort.,  1865,  15:  293-295. 

.     1866.     Production  de  varietes  par  le  bouturage.     Hort.  Frangais,  8:  237-240. 

.     1868.     Coleus  nouveaux  des  Anglais.     Hort.  Frangais,  10:  271-274. 

Hooker,  W.  J.     1853.    Coleus  blumei.   Curtis'sBot.  Mag.,  iii, 9: pi.  4754  (with description). 
Hub,  Henri.     1914.     The  origin  of  X  Capsella  bursa-pastoris  arachnoidea.     Am.  Nat.,  48: 

193-235. 
Keeble,  Frederick,  and  E.  F.  Armstrong.     1912.     The  role  of  oxydases  in  the  formation 

of  the  anthoeyan  pigments  of  plants.     Jour.  Genetics,  2:  277-311,  pi.  19. 
KtJSTER,    Ernst.     1913.     Ueber    Zonenbildung  in    KoUoidalen   Medien.     Beitrage   zur 

entwicklungs  mechanischen  Anatomie  der  Pflanzen. 
Lemoine,  Ch.     1861.     Coleus  verschaffeltii.     111.  Hort.,  8:  pi.  293  (with  description). 
Lindemuth,  H.     1897.     Vorlaufige  Mitteilungen  von  Veredelungs  versuchen  innerhalb  der 

Malvaceen  und  Solanaceen.     Gartenflora,  46:  1-6. 
.     1899.     Kataibelia  vitifolia  (Willd.)  mit  goldgelb  marmorierten  Blattern.     Gar- 
tenflora, 48:  431-434. 

.     1901.     Impfversuche  an  Malvaceen.     Gartenflora,  50:  8-11. 

.     1905.     Ueber  verschiedene  Arten  der  Panaschiire,  deren  Uebertragbarkeit  durch 

Transplantation  und  Samenbestandigkeit.     Gartenflora,  54:  125-128. 
.     1907.     Studien  iiber  die   sogenannte  Panaschiire  und    iiber  einige  begleitende 

Erscheinungen.     Landw.  Jahrb.,  36:  807-862. 
Mendel,  Gregor.    1865.    Versuche  iiber  Pflanzen-Hybriden.    Verhandlungen  des  Natur- 

forschendenvereines  in  Briin,  4:  (Abhandlungen)  1-47.     (Translation  in 

"Mendel's  Principles  of  Heredity"  (third  impression),  1913,  335-379.) 
Moore,  Thomas.     1868.     New  hybrids  of  Coleus.     Gard.  Chron.,  33:  376-377. 

.     1869.     Coleus  Queen  Victoria.     Florist  and  Pomologist,  1869,  1-2,  pi. 

Morgan,  T.  H.     1913.     Factors  and  unit  characters  in  Mendehan  heredity.     Am.  Nat.,  47: 

.5-16. 
Morren,  Edouard.     1856.     Le  Coleus  blumei,  variete  pectinatus.     Belg.  Hort.,  6:  99,  pi. 
Nilsson-Ehle,  H.     1908.     Nogat  om  nuvarande  principer  vid  hosthveteforadhngen  pa 

Svalof.     Sveriges  Utsadesforenings  Tidskrift.  ^ 
.     1909.     Kreuzungsuntersuchungen  an  Hafer  und  Weizen.     Lands  Univ.  Arssk., 

Teil.  1. 
Overton,  E.     1899.     Beobachtungen  und  Versuche  iiber  das  Auftreten  von  rothem  Zell- 

saft  bei  Pflanzen.     Jahrb.  Wiss.  Bot.,  33:  171-231. 
Planchon,  J.  E.     1852.     Coleus  blumei.     Fl.  Serres,  8:  141. 
Pitnnett,  R.  C.     1911.     MendeUsm.     Third  edition. 

Pynaert,  Ed.     1881.     Coleus  reine  des  Beiges.     111.  Hort.,  28:  102-104.     pi.  425. 
Reid,  Katherine  W.     1914.     Variegated  abutilons.     Jour.  N.  Y.  Bot.  Gard.,  15:  207-213. 
Riddle,  Oscar.     1909.     Our  knowledge  of  melanin  color  formation  and  its  bearing  on 

Mendehan  description  of  heredity.     Biol.  Bull.,  16:  316-351. 
RoDiGAS,  Em.     Coleus  blumei  Benth.  var.  nov.  Coleus  Gloire  de  Dijon.     III.  Hort.,  35:  31, 

pi.  46. 

.     1892.     CoZeas  Wwwei  Benth.  varietates  novae.     111.  Hort.,  39:  111-112,  pi.  164. 

Shull,  G.  H.     1908.     The  pedigree  culture:  its  aims  and  methods.     Plant  World,   11: 

21-28,  55-64. 
.     1911.     Defective  inheritance  ratios  in  Bursa  hybrids.     Verh.  Nat.  Ver  Brunn, 

49:  1-12. 

.     1912.     "Phenotype"  and  "clone."     Science,  ii,  35:  182,  183. 

.     1914.     Ueber  die  Vererbung  der  Blattfarbe  bei  Melandrium,.     Ber.  Deutsch.  Bot. 

Gesell.,  31:  (40)-(80). 
Stout,  A.  B.     1913.     A  case  of  bud-variation  in  Pelargonium.     Bull.  Torrey  Club,  40: 

367-372,  pi.  20. 
Verlot,  J.  B.     1866.     Plantes  nouvelles,  rares  ou  peu  connues.    Revue  Hort.,  37:  279-280. 
Wakeman,  Nellie.     1911.     The  Monardas.     Bull.  Univ.  Wis.  Sci.,  4:  81-128. 
Webber,  H.  J.     1903.     New  horticultural  and  agricultural  terms.     Science,  ii,  18:  501- 

503. 
Wheldale,  M.     1914.     Our  present  knowledge  of  the  chemistry  of  the  Mendehan  factors 

for  flower-colour.     Jour.  Genetics,  4:  109-129.     pi.  7. 
Witte,  H.     1862.     Coleus  verschaffeltii  Lem.  et  son  introduction  en  Em-ope.     Ann.  Hort. 

Bot.,  5:  12.5-128. 
Woods,  A.  F.     1899.     The  destruction  of  chlorophyll  by  oxidizing  enzymes.    Centralb. 

Bakt.  II,  Abt.  5:  745-754. 


BY   THE    SELECTION    OF    SOMATIC    VARIATIONS.  79 

EXPLANATION  OF  FIGURES  IN  PLATES  1.  2.  3.  AND  4. 

The  paintings  here  reproduced  were  made  by  Misa  Mary  Eaton,  artist  of  the  New  York 
Botanical  Garden.     All  figures  are  reduced  to  about  three-fourths  natural  size. 

Plate  1. 

Fig.     1.     A  typical  leaf  of  the  pattern  yellow-red  blotched  taken  from  plant  1171013  on 
February  21,  1914. 
2.     The  pattern  green-yellow-red  blotched.     Taken  from  plant  121  on  April  1.1,  1912. 
The  yellow  border  is  somewhat  irregular  and  is  not  so  fully  developed  as  in 
summer. 

4.  Leaf  classed  as  green-yellow  spotted-red  blotched.     Painted  on  January  .30,  191.3. 

Shows  rather  few  scattered  yellow  spots  and  a  few  rather  large  epidermal 
blotches. 

5.  Pattern  green-red  blotched  from  plant  131,  taken  April  1.5,  1912.     Shows  the  com- 

plete loss  of  yellow  as  it  occurred  in  the  first  bud  variation  observed  and  as  it 
frequently  appears  in  plants  having  yellow. 
5a.  Young  leaf  of  the  pattern  green-red  blotched. 

6.  The  pattern  yellow-green-red  blotched.     Taken  from  plant  1171  on  November  2, 

1912.  Shows  the  type  which  appeared  on  a  plant  with  the  pattern  of  figure  2 
by  a  reversal  of  the  relative  positions  of  the  green  and  yellow. 

7.  A  good  example  of  the  laciniate  type  of  leaf.     Taken  on  January  31,  1912,  from 

a  plant  quite  identical  to  plant  123153  shown  in  plate  4. 

8.  The  green-yellow-solid  red  type.     Differs  from  figure  2  in  having  a  solid  red  instead 

of  a  red  blotched  epidermis.  Taken  on  January  4,  1913,  from  plant  32  and 
shows  the  typical  development  of  the  yellow,  which  was  very  constant  and 
uniform  on  the  plant  diu-ing  the  winter. 

9.  A  typical  leaf  of  the  green-solid  red  type.     This  differs  from  figure  8  only  in  having 

no  yellow  in  the  subepidermal  tissues  and  from  figure  .5  in  possessing  a  solid 
red  epidermis. 
9a.  A  young  leaf  of  the  pattern  green-solid  red.     The  red  completely  covers  the  leaf 
and  is  of  the  same  intensity  as  in  a  mature  leaf. 

10.  This  figure  shows  the  upper  surface  of  a  leaf  of  the  i^attern  green-yellow-solid  red 

upper  center.  Figure  lOa  is  of  the  under  surface  of  the  same  leaf.  The  dis- 
tribution of  the  green  and  yellow  is  as  in  figure  2.  The  red  pigmentation  is 
almost  entirely  massed  in  the  epidermis  of  the  upper  surface.  There  are  a 
few  small  areas  of  red  in  the  lower  epidermis.     Painted  on  December  9,  1913. 

1 1 .  The  type  yelloiv-green-solid  red  painted  on  October  26,  1914.     This  tj^pe  developed 

from  type  of  figure  8  by  a  reversal  of  the  relati\'o  positions  of  the  green  and 
yellow,  a  change  which  also  gave  figure  6  from  figure  2. 

Plate  2. 

Fig.  12.  Type  green-yellow,  painted  May  17,  1914.  The  i)attein  is  slightly  irregular  in 
this  leaf,  with  a  yellow  segment  extending  to  the  midrib.  This  pattern  was 
derived  from  type  of  figure  2  by  a  loss  of  epidermal  red.  There  are  a  few 
streaks  of  red  in  the  vascular  tissues. 
13.  The  green  pattern  showing  some  epidermal  red  in  the  vascular  tissues  and  a  pale 
and  diffuse  shade  of  yellow.     Painted  on  June  5,  191  4. 

13,  a,  b,  c,  d.     Four  successive  leaves  of  a  plant  of  the  same  line  of  descent  as  the  plant 

of  figme  13.  Painted  on  February  2,  191.5.  The  series  shows  an  increase  of 
red  in  the  subepidermal  tissues  as  the  leaf  matures,  with  tlie  maxinunn  develop- 
ment in  figure  d.  Comparison  with  figure  13  shows  the  increase  of  red  in  a 
hne  of  descent  by  gradual  fluctuation.  The  distribution  of  red  indicates  a 
relation  involving  vascular  tissues. 

14.  Pattern  yellow-green.     This  was  derived  from  type  G  by  a  loss  of  epidermal  red. 

The  streaks  of  red  are  subepidermal.     Painted  on  February  11,  1914,  showing 
the  decreased  development  of  yellow  frequent  in  this  type  during  winter. 
14a.  The  typical  summer  condition  of  the  type  yellow-green.     Painted  on  May  29, 
1914.     The  yellow  areas  are  bounded  in  marked  degree  by  veins  and  there  is  a 
much  less  development  of  green  cells  in  the  central  area. 


80  ESTABLISHMENT    OF   VARIETIES    IN    COLEUS. 

Plate  2 — Continued. 

Fig.  15.  A  leaf  with  no  yellow  and  with  almost  no  red  on  the  under  surface.  A  type  quite 
like  that  described  for  the  original  Coleus  bluniei.  This  type  appeared  thus 
far  only  as  a  fluctuating  variation  as  a  winter  condition  of  a  few  plants  of  the 
type  of  figure  10.  At  the  time  this  was  painted,  January  24,  1914,  the  oldest 
leaves  of  the  plant  were  typical  for  type  10  and  the  younger  leaves  were  uni- 
form for  the  coloration  here  shown.  The  under  surface  of  this  leaf  had  only 
a  few  red  blotches  similar  to  those  of  figure  lOo. 

17.  A  leaf  with  irregular  pattern,  developed  from  type  of  figure  2. 

18.  One  of  the  leaves  of  the  plant  shown  in  figure  21,  showing  the  absence  of  yellow  in 

half  of  a  leaf. 

19.  Leaf  painted  on  January  18,  1914,  showing  rather  marked  increase  of  yeUow  in  the 

type  of  figure  2  during  winter.  Painting  was  made  after  the  yellow  areas  had 
begun  to  turn  to  white. 

20.  Painted  on  July  5,  1913.     Typical  of  the  most  extreme  fluctuation  of  type  figure  2 

in  regard  to  increase  of  yellow. 

Plate  3. 

Fig.  21.  Young  plant  12514,  grown  from  a  bud  showing  sectorial  loss  of  yellow,  by  bud 
variation  in  half  of  the  bud.     Photographed  and  painted  on  November  9,  1912. 

22.  One  of  the  leaves  in  a  branch  showing  sectorial  loss  of  green  in  the  pattern  of 

figure  6.  In  this  leaf  the  loss  appeared  in  one  side  of  the  leaf.  Painted  on 
December  4,  1913. 

23.  Leaf  classed  as  green-yellow  spotted-red  blotched.     The  red  epidermal  blotches  are 

large  and  much  coalesced,  which  is  a  frequent  variation  from  the  condition  of 
figure  5. 
24  and  24c.  Lower  and  upper  surfaces  of  the  same  leaf  from  a  young  plant  grown 
from  a  bud  variation  which  involved  a  sectorial  loss  of  red,  giving  solid  red 
upper  center  from  solid  red  on  both  surfaces.  In  this  leaf  these  two  types  are 
sharply  limited  to  one  half  of  the  leaf.  The  young  plant  exhibited  loss  of 
yellow  by  fluctuating  variation.  The  leaf  painted  was  an  upper  leaf  showing 
no  yellow,  but  at  the  same  time  the  older  basal  leaves  possessed  much  yellow, 
as  in  type  of  figure  10. 

25.  Leaf  painted  on  March  25,  1913,  showing  sudden  bud  variation  affecting  only  the 

half  of  a  leaf.  The  plant  possessed  leaves  of  the  type  figure  2  shown  in  the 
left  side.  A  reversal  of  the  relative  position  of  the  green  and  the  yellow  gave 
the  pattern  of  figure  6  in  the  right  side  of  the  leaf,  as  shown. 

26.  Taken  on  March  28,  1913,  showing  the  fluctuational  decrease  of  yellow,  giving  a 

poorly  defined  yellow  border.  During  the  f  ollowmg  summer  the  plant  returned 
to  the  typical  form  of  type  2.  This  leaf  is  quite  typical  of  the  increase  of  green 
during  winter  seen  in  numerous  plants  of  type  2,  as  mentioned  on  page  22  of 
the  text. 

27.  Painted  on  July  27,  1913,  showing  an  in-egular  pattern  with  yellow  at  the  border 

about  the  apex  of  the  leaf  and  illustrating  a  fluctuation  produced  from  both 
types  2  and  4.     Leaf  also  shows  irregular  distribution  of  epidermal  red. 

28.  Leaf  classed  as  green-red  blotched,  but  with  few  large  blotches  somewhat  run 

together. 

29.  Leaf  also  classed  as  green-red  blotched,  but  with  fine  blotches  much  coalesced. 

Plate  4. 

Three  plants  photographed  on  January  16,  1914,  all  descended  from  branch  2  of  plant  1. 
Plant  125111  has  leaves  uniformly  entire  and  of  the  pattern  green-yellow-red  blotched  (fig.  2). 

Tips  pinched  off  to  prevent  early  flowering  in  greenhouse. 
Plant  1251412  has  leaves  uniformly  entire  and  of  the  pattern  green-red  blotched. 
Plant  123153.     Old  leaves  entire,  youngest  leaves  deeply  laciniate.     Typical  condition 
during  early  winter  for  plants  showing  the  fluctuation  in  leaf  shape. 


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